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Full text of "Bulletin of the British Museum (Natural History), Geology"

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BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



]W 



rS'. 



GEOLOGY 
VOL. XI 

1965— 1966 



TRUSTEES OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

LONDON: 1967 



DATES OF PUBLICATION OF THE PARTS 

No. I 

No. 2 

No. 3 

No. 4 

No. 5 

No. 6 

No. 7 

No. 8 

No. 9 
No. lo 



Qth July 










1965 


7th September 










1965 


8th December 










1965 


8th December 










1965 


17th December 










1965 


7th January 










1966 


nth January 










1966 


2 1st January 










1966 


28th January 










1966 


19th April . 










1966 



PRINTED IN GREAT BRITAIN 
BY ADLARD & SON LIMITED 
BARTHOLOMEW PRESS, DORKING 



CONTENTS 

GEOLOGY VOLUME XI 

PAGE 

No. I. The head of Dipterns valenciennesi Sedgwick & Murchison. 

E. I. White i 

No. 2. Some Lower Cretaceous Terebratelloidea. E. F. Owen 47 

No. 3. Middle Jurassic Ostracoda from the Grey Limestone Series, 

Yorkshire. R. H. Bate 73 

No. 4. Human skeletal material from Ceylon, with an analysis of the Island's 

Prehistoric and Contemporary Populations. K. A. R. Kennedy 135 

No. 5. Some new British Albian Ostracoda. P. Kaye 215 

No. 6. Tertiary Red Algae from Borneo. J. Harlan Johnson 255 

No. 7. British Wealden Sharks. C. Patterson 281 

No. 8. On certain Triassic and Liassic representati^'es of the family 

Pholidophoridae s. str. O. Nybelin 351 

No. 9. Some British Jurassic and Cretaceous Ostracoda. F. W. Anderson 

& D. Barker 433 

No. 10. Trilobites of the Henllan Ash, Arenig Series, Merioneth. H. B. 

Whittington 489 

Index to Volume XI 507 



THE HEAD OF 
DIPTERUS VALENCIENNESI 

Sedgwick & Murchison 




ERROL IVOR WHITE 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

GEOLOGY Vol. ii No. i 

LONDON: 1965 



THE HEAD OF 
DIPTERUS VALENCIENNESI 

Sedgwick & Murchison 



BY 

ERROL IVOR WHITE, F.R.S. 




Pp. 1-45 ; 3 Plates ; 51 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. 11 No. i 

LONDON: 1965 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of larger 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 1 of the Geological 
[Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 



Trustees of the British Museum (Natural History) 1965 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued gth July, 1965 Price Twenty-five Shillings 



THE HEAD OF 
DIPTERUS VALENCIENNESI 

Sedgwick & Murchison 
By ERROL IVOR WHITE 

SYNOPSIS 

There is only one known species of Dipterus from the Middle Old Red Sandstone of Scotland, 
D. valenciennesi Sedgwick & Murchison. The split between the dipnoan and crossopterygian 
stocks took place at the mosaic stage of evolution of the skull-roof bones, and the nomenclature 
of the consequent roofing-plates in one group has no relevance to that in the other ; entirely 
independent systems must be used. A modified version of Forster-Cooper's alphabetical system 
is accepted for the dipnoans, the pattern being developed by invasion and loss rather than by 
fusion. There has been a gradual loss of rows of plates at the back of the dipnoan head with 
consequent movement forward of the occipital cross-commissure. The details of the neurocranium 
are described. 

The Name 

Specimens of Dipterus from the Middle Old Red Sandstone of Scotland have generally 
been referred to a single species, Dipterus valenciennesi Sedgwick & Murchison, al- 
though from time to time attempts have been made (e.g. Pander 1858 : 7 ; Watson & 
Day 1916 : 29) to separate the solid-snouted forms under Agassiz's (1844 : 29) specific 
name of platycephalus. More recently Westoll (1949 : 127-128) decided that the 
Banniskirk specimens showed sufficient peculiarities in the skull-roof pattern to 
warrant their separation as a distinct species, and in the belief that all Sedgwick & 
Murchison's (1829 : 143, pi. 15, figs. 1-3, pi. 16, figs, i, 3, pi. 17, figs. 1-3) original 
specimens of their three species, D. brachypygopterus, D. macropygoptenis and D. 
valenciennesi came from Banniskirk quarry, has used the first of these names, 
D. brachypygopterus, for the Banniskirk form which he considered distinct, dropping 
the names D. macropygopterus and D. valenciennesi altogether as synonyms of D. 
brachypygopterus , and has revived Agassiz's name of platycephalus for all the other 
Scotch specimens of Dipterus, with or without solid snouts. 

Unfortunately the provenance of all Sedgwick & Murchison's figured material, now 
in the collection of the Geological Survey, is not so certain. Only one of the specimens, 
a co-type of Dipterus macropygopterus (pi. 15, fig. 2 ; G.S. no. 6448) was originally 
labelled "Banniskirk", all the others being labelled " Thurso ". " Thurso " indeed 
was used in early days to cover Banniskirk, which is some 8 miles to the south-east 
of the town, but it also covered many other quarries in the neighbourhood, and 
identification of the locality must depend on the recognition of matrix, a very 
doubtful procedure in these variable rocks. Sedgwick & Murchison's remark about 
" those from Banniskirk" (1829 : 142) refers to the " small number (sent to Cuvier) 
of the whole series afterwards examined bj' Messrs. Valenciennes and Pentland " 
(p. 142, footnote), and indeed on the previous page Sedgwick & Murchison are at 
pains to point out that " when the attention of geologists was first drawn to these 
ichthyolites, it was not known that specimens of them were to be found in any other 



THE HEAD OF DIPTERUS V A LE N C I E N N E S I 



quarry than that of Banniskirk. The authors of this memoir have however, since 
discovered that similar remains are extensively . . . spread over the Caithness deposit, 
etc. etc. . . ." This refers to Old Red fishes in general, but clearly puts to question 
the origin of the specimens they figured, which are from the" more perfect remains" 
sent to Valenciennes & Pentland. 

But even more important is the lack of substance in the supposition that the 
Banniskirk forms have a special pattern of their head plates, qualifying them for 
specific recognition. None of Sedgwick & Murchison's figured specimens shows the 
head-pattern clearly, and so are irrelevant to this point, whether they come from 
Banniskirk or not. All the five specimens from Banniskirk figured by WestoU (1949 : 





Figs, i, 2. Dipterus valenciennesi S. & M. Skull-roof showing normal pattern. Bannis- 
kirk, Caithness. Fig. i, Geol. Surv. No. 6464. Fig. 2, R.S.M. 1859.33.623 (Hugh Miller 
Coll.). Both X3. 



137-139. text-fig. 4) show to some degree brachypygopierus features, that is, the 
paired bones C are replaced by the forward extension of B, and to a small extent by 
the medial growth of some of the lateral series K-M, although he notes that it is 
possible that something very hke the Achanarras type (" D. platycephalits ") occurs 
very rarely at Banniskirk. 

One such individual was figured by Pander (1858, pi. i, fig. 2), and his diagram 
(pi. 3, fig. i) is essentially " normal " in pattern (Text-fig. 10). Of the six specimens 
available to me that bear original Banniskirk labels and show the roof pattern 
clearly, four (G.S. 6464, R.S.M. 1859.33.623, and R.S.M. 1876. 18.31; Text-figs. 
1-3) and R.S.M. 1859.33.6261 show the typical normal pattern with 2 " C " plates 

1 The initials R.S.M. indicate that the specimen came from the Royal Scottish Museum ; D.M.S.W. 
from Professor Watson's collection ; G.S. from the Geological Survey ; and M.M. from Manchester 
Museum. Other specimens numbered with or without an initial P, are in the British Museum (Natural 
History). 



THE HEAD OF DIPTERUS V A LE N C I EN N E S I 5 

and a median " D ", as does a fifth (33149I, Text-fig. 4) whose supposed provenance 
from Banniskirk has been subsequently added in pencil, presumably on the basis of 
the matrix. None of these five specimens would have caUed for comment in regard 
to its skull-roof. 

However, on a slab recently collected at Banniskirk by Mr. Jack Saxon is a small 
nearly complete fish whose head certainly does call for comment, but for another 
reason (P.46762D, Text-fig. 8). Two C's, a D and 2 E's are well developed in the 
normal way, but the hinder part of the skull is completely asymmetrical with no J 
on the left side, its space being largely occupied by B much swoUen in front, but with 
some help from Lj and K, on to which the rather unusual anterior pit-line extends to 
meet the supraorbital sensory canal. There is no indication of fusion here whatever, 
and indeed it is most likely that this arrangement is due to the invasion by B, and to a 
lesser extent by L^ and K, of the area where J has failed to develop, in accordance 
with the sensible principle laid down by Parrington (1950 : 537-540) that fusion 
should not be postulated unless clearly indicated. 

Only two of the six specimens with original Banniskirk labels show the " brachy- 
pygopterus " pattern with the greatly enlarged plate B and no C's (42480, R.S.M. 
1859.33.624, Text-figs. 5, 6), and both these specimens seem to indicate by the 
structure of plate B, which in each shows radiation from a single centre, that in these 
specimens also we have instances of plates (B) invading areas where the growth of the 
normal plates has been inhibited. But that it was not always entirely a matter of 
simple invasion is shown by one of Peach's specimens, a small skull-roof collected in 
1858 at Sandside, some 10 miles west of Thurso. In this (R.S.M. 1887.35, Text-fig. 7) 
B is much enlarged forwards and the L bones, two on the left and one on the right side, 
are wide; but between their anterior ends are two very small C plates, partly fused 
together and partly fused to B. The front part of the skull, well separated by a post 
mortem displacement from the hinder part, shows a small M on the left, a larger M on 
the right, large N's on both sides, a wide single E incompletely separated from the 
snout, and behind this a wide plate which is evidently compound. Whether this last 
represents D+ is not very important, but the definite natural break behind the well- 
defined posterior border does seem to establish that it was not part of C. 

A reasonable and simple explanation of the form of the B and C and L bones in this 
specimen is provided by Parrington's (1956 : 395-404) " Patterns of dermal bones ": 
the development of the C bones has been retarded and much of their space occupied 
by B and the L bones before its growth began, and when it did make progress 
growth-pressure to the rear induced partial fusion with B and lateral pressure induced 
fusion between the two C's. That neither C bone tended to fuse with its lateral 
neighbour L, is noteworthy, for Westoll (1949 : 135) has already remarked on the 
reluctance of " general " bones to fuse with " lateral line bones ", and figured only 
two possible examples of this (1949 : text-figs. zB, ^E). 

The " brachypygopterus " pattern is not a specific character but an occasional 
variation, not uncommon at Banniskirk but occurring infrequently elsewhere, as at 
Achanarras (Parrington 1950 : 540, text-figs. 4, 5). Most of the specimens showing 
this pattern are smaU, but not all, witness Parrington's second specimen. 

So far as present knowledge goes one is dealing with a single variable species, and 
for this all three of Sedgwick & Murchison's specific names of 1828 are available (the 



THE HEAD OF DIPTERUS V ALEN C I EN N E S I 



fourth name macrolepidotus refers to an osteolepid, Thursius — see Woodward 1891 : 
373) ; but Agassiz's platycephalns of 1844 is not (Text-fig. 8). And Agassiz (1835 : 115) 
having used the inadmissible name " macrolepidotus ", Pander (1858 : 6) as the first 
acceptable reviser, had the right to select whichever of the three available names that 
he wished for the specific name (Article 24, Internat. Code Zool. Nomenclature, XV 

3 





V-- 



FiGS. 3-5. Dipterus valenciennesi S. & M. Fig. 3, Skull-roof showing normal pattern. 
Banniskirk, Caithness. R.S.M. 1876. 18.31. X3. Fig. 4, Skull-roof showing normal 
pattern. PBanniskirk. 33149(1). X3. Fig. 5, Skull-roof showing " brach5rpygop- 
terus " pattern. Banniskirk. 42480 (Peach Coll.). X2-25. 



Internat. Congr. Zool. 1961) : and his choice of " valenciennesi " was fully supported 
by Woodward (1891 : 236). So the name of the species remains Dipterus valencien- 
nesi Sedgwick & Murchison. The term " brachypygopterus-type " may be used for 
the form of D. valenciennesi described by WestoU from Banniskirk, but from any 
locality, in the way that platycephalus-type has been used for the solid-snouted 
forms. In regard to this last form the original head-shield, the lectotype, upon 
which Agassiz based his " Polyphractus platycephalus " is unusually wide and smaU, 
for the numerous resorption lines and large openings of the foramina seem to indicate 



THE HEAD OF DIPTERUS V A LE N C I EN N E S I 7 

an aged fish (P. 3373«, Text-fig. 51). Its small adult size, a little over 4 cm. in length, 
is rather unusual in the " platycephalus-type ". Forster-Cooper (1937 : 231) 
noticed a general correlation between size and the development of the snout, and it 
seems true that while the hard snout can occur occasionally in rather small specimens, 
such as this and the original of Text-fig. 7, it is invariably present in large speci- 
mens. This does not support Marshall's (1962 : 314) remark that the difference 
between the soft and hard snouted forms" may be explained as seasonal differ- 
ences " : they seem more likely to be linked with size and age. 

It is difficult to give absolute measurements of breadth and length in these skulls, 
but in general the larger skulls tend to be relatively narrower, up to about 15%, and 
while the smaller specimens, those measuring under 6-o cm. from the back of B to the 
tip of the hard snout, have only 2 or 3 plates along the sensory canal series as a rule, 
i.e. Y2-N, not all the larger specimens show multiplication. But there are obvious 
exceptions to both these tendencies. 

Nomenclature of the Bones of the Skull-roof 

From the first the identification of the dermal bones of the skull-roof in lung-fishes 
gave rise to much difficulty, in the fossil forms because they had too many plates for 
exact correlation with those of the general run of fishes, and in the living because they 
had too few ; and usually the earlier zoologists, such as Giinther (1871), Huxley 
(1876) and Bridge (1898) wisely treated them with caution. The first serious attempt 
to identify the plates of the head of the fossil forms according to the nomenclature in 
general use for vertebrates was made by D. M. S. Watson (Watson & Day 1916 : 29- 
32 ; Watson & Gill 1923), but his interpretation met with flat contradiction from 
Goodrich (1930 : 305). 

Since then efforts to bring order into the labeUing of dipnoan roofing bones has 
developed along two very divergent lines : one a purely arbitrary alphabetical 
notation, proposed by Forster-Cooper (1936) : and the other the compound nomen- 
clature of Holmgren & Stensio (1936), which seems in effect a more or less spatial use 
of the general vertebrate terms based on the assumption of widespread fusion of 
plates. 

In a recent study of the skull-roof of a dipterine from Belgium (White 1962) the 
alphabetical notation of Forster-Cooper (1937 : 228-229) was used without conviction 
or enthusiasm, merely because the alternatives seemed even less desirable. Romer's 
(1936 : 242, text-fig. 9) code-system, if more logical in intention, is, as Westoll 
(1949 : 126) has pointed out, incorrect in application. 

On the other hand Lehman's (1959 : 6) advocacy of Holmgren & Stensio's (1936 : 
365, text-figs. 280 A~C) terminology, based on the theory of widespread fusion, is 
only acceptable if the homologies of the component bones are generally agreed, which 
they certainly are not. 

Jarvik (1948 : 81, 291), while strongly supporting the fusionists, does admit that 
complex bones in different groups of fishes given the same name, rarely, if ever, are 
strictly homologous, " even in closely allied groups ". He further remarks (p. 83) that 
" It is true that dermal bones are variable in shape and sometimes may invade the 
territories of adjoining bones. Thus it seems likely that the individual variations in 



THE HEAD OF DIPTERUS V A LE N C I E N N E S I 





Figs. 6-8. Dipterns valenciennesi S. & M. Fig. 6, Skull-roof, with left cheek, showing 
" brachypygopterus " pattern. BanniskLrk, Caithness. R.S.M. 1859.33.624 (Hugh 
Miller Coll.). X3. Fig. 7, Skull-roof showing pattern intermediate between normal 
and "brachypygopterus" types. Sandside, Caithness. R.S.M. 1887.35 (Peach Coll.). 
X3- Fig- 8, Skull-roof of very small fish showing extremely asymmetrical hinder end. 
Banniskirk. P.46762D (Colld. J. Saxon, 1964). X3-5. 



THE HEAD OF DIPTERUS VALENCIENNESI g 

shape and extent of the dermal bones of the cranial roof in Osteolepi formes . . . are 
not always to be explained by various fusions, etc. ", which comes quite near to 
WestoU's (1944 : 114) assertion that " the compound form (of nomenclature) . . . 
expresses no more than geographical extent . . .", and certainly weakens the argu- 
ment for too rigid adherence to the hypothesis of universal fusion without unduly 
encouraging those who pin their faith overmuch on the apparently contradictory 
idea of "loss and invasion " — " aut Caesar aut nihil " may be a good political precept 
but it is usually a poor biological principle — indeed, Truth in Nature has a habit of 
falling between two schools of thought. 

In spite of what has been said by their proponents concerning the advantages of 
either philosophy for establishing the identity or homologies of bones or their supposed 
components in Dipnoi, it is in practice often very difficult to identify bones by any 
system satisfactorily, and important mistakes and man}^ doubtful identifications 
have been made on both sides. Not the smallest source of error has been due to 
attempts to establish the course of the canals by the openings of the tubuli on the 
surface of the bones; and this in the nature of things is often inevitable, since it is not 
always desirable or possible to dissect out the canals. 





Figs. 9, 10. Diagrams of skull-roof of Dipterus. Fig. 9, Diagram showing Romer's (1936 : 
243, text-fig. \c) alphabetical notation. Fig. 10, Pander's diagram of the skull-roof 
(1858 : 55, pi. 3,'^fig. I). 



lo THE HEAD OF DIPTERUS V A LE N C I E N N E S I 

Forster-Cooper's alphabetical notation (FCAN) 

Forster-Cooper's {1937 : 228, text-fig. 3) alphabetical notation (FCAN) was based 
on a very simple plan, but unfortunately he never developed it fuUy, and sometimes 
used it incorrectly {e.g. pi. 7, figs. 13, 14). 

It should be noted that this scheme was quite arbitrary and not based on the 
sensory canals and pit-hnes : indeed, it is quite plain that Forster-Cooper had no 
clear idea of their true course. In almost every specimen that he figured and labelled 
he relied on the external openings of the tubuli of the sensory canals to determine 
their position, and sometimes mistook damaged tubercles of the ornament and post 
mortem cracks for pores and pit-lines (1937 : 233, pi. 5, fig. 9, text-fig. 5). 

His description of the lateral lines is indeed most puzzling : " The course of the 
main lateral line from the body on to the head appears to run across the supra- 
cleithrum to the bone H. It then runs to I, where it branches, one branch going 
through Y, the other to B, where it meets its feUow of the opposite side. This junction 
is more noticeable in the platycephalus condition than in that of valenciennesi. Just in 
front of this point there is a V-shaped pit line pointing forwards whose arms run 
through the paired J bones from each side ". 

In the next paragraph he stated that " The pores marking the underlying sense- 
organ canals always occur on the same series of bones in a line running from G through 
H, Y, X, K, L, O, P, Q, and then round the front of the snout at the anterior edges of 
EandF ". 

Not only are these two statements contradictory, but both are wrong. For con- 
venience they have been translated on to Forster-Cooper's original diagram (Text- 
fig. 11), the first statement shown by solid dots on the right side, the second by hoUow 
dots on the left. Neither is supported by any of his own touched-up photographic 
illustrations, including that of his standard specimen (Forster-Cooper 1937 : 228, pi. 
4, fig. 7, text-fig. 3) on which his diagram was based. 

Apparently these statements were based on Goodrich's (1925 : ^^, text-figs, i, 3, 4) 
descriptions and figures, which do place the occipital commissure across Goodrich's 
" tabular " (H), " Post-parietal " (I) and " dermal supraoccipital " (B) just behind 
the anterior pit-hnes, with some indication in Dipterus of a forward branch in the 
" post-parietal ", while the " supra-temporal " and " supraorbital " canals run in a 
continuous sinuous curve very much as Forster-Cooper describes. It may be noted 
that the much earlier diagram of that very acute observer C. M. Pander (1858 : 55, 
pi. 3, fig. i), is a perfectly accurate representation of the skull-roof of Dipterus and 
its sensory canals (Text-fig. 10). 

The general course of the canals and pit-lines is now well known, thanks to the 
work of Graham-Smith & WestoU (1937 : 244, text-fig. 2a), WestoU (1949 : 126, text- 
figs. 2, 3 etc.) and Parrington (1950 : 536, text-fig. i). In the first paper Graham-Smith 
& WestoU partly rationalized Forster-Cooper's notation, using a rather better pre- 
served head, P . 22189 from Achanarras, as the standard specimen ; plate G, which in 
Forster-Cooper's specimen (see Text-figs. 11, 12, 21) was a body-scale unconnected 
with the lateral line, was dropped; Y was allotted to two plates, and so was L on the 
right side as the normal pattern, the single plate on the left being considered the 
result of fusion. For no apparent reason Q was substituted for P. However, 
WestoU's remarks (1949 : 130-134) that " All these latero-sensory canal bones are 



THE HEAD OF DIPTERUS V A L E N C I E N N E S I 




Figs. 11-13. Dipterus valenciennesi S. & M. Fig. 11, Forster-Cooper's " Generalized 
diagram " (1937 • 228, text-fig. 3) with the sensory canals and pits added from his 
description (p. 233), the solid dots on the right side show " the course of the main lateral 
line " ; the rings on the left, the occurrence of the " pores marking the underlying sense- 
organ canals. " Fig. 12, Sketch of the specimen on which Fig. 11 was based showing the 
distribution of the pores of the sensory canals (and of the canals themselves on the 
partly dissected plates X, J, K, L, N, P) and of the pit-lines. Achanarras, Caithness. 
P. 22187. X2-3. Fig. 13, Corrected diagram of the same specimen with emended 
lettering. 



12 THE HEAD OF DIPTERUS V A L E N C I E N N E S I 

extremely constant in their development, with the exception . . . that adjacent 
members 'fuse' ", and " There is never any difficulty in deciding where 'fusion' has 
occurred at least in the skull-roof " seems to the present writer to be very optimis- 
tic. Indeed WestoU's figures (1949, text-figs. 2, 3 etc.), like the earher figures of 
Stensio, are diagrammatic sketches, and the supposed source of the sensory canals was 
similarly based on the external pores. 

Lack of more precise information doubtless accounts for WestoU's (1949 : 136) 
remark that " these canals show no abnormalities in the specimens studied ". 

Dissections have been carried out in a number of specimens, including both Forster- 
Cooper's and Graham Smith & WestoU's standard specimens with interesting results. 

In Forster-Cooper's specimen (P. 22187) part of the left side of the head has been 
prepared (Text-fig. 12) and the pattern is shown in diagrammatic form (Text-fig. 13). 
In the latter it will be noted that there is some alteration of the original lettering. G, 
which has already been noted as a scale, is omitted ; H, since it belongs to the main 
sensory canal series with Y and X, is changed to Z. But of much more significance is 
the absence of Lj^ and M and the labeUing of Y as Yj^ -f Yg.^ Here we are brought up 
against the main problem of fusion and/or replacement. 

West oil (1949 : 126, 128) although keenly critical of the fusionist tendencies of 
Stensio and his school in relation to large bones, nevertheless stated that " Onto- 
genetical studies on hving bony fishes such as Amia show that true fusion of adjacent 
bone rudiments developed along a latero-sensory canal is a normal feature of the 
development of large bones. It is here accepted that fusion of this type occurs in 
Dipterus " and gives (1949 : 132) some 45 instances where fusion of lateral-line 
elements are supposed to have occurred. 

Positive detection of such fusion in more or less adult fossil skuUs would presumably 
depend on how rudimentary the bone-rudiments were at the time of fusion : complete 
fusion of the earliest stages would not probably be traceable, and whether it took 
place or not must depend on other indirect evidence, if any. 

Parrington (1950 : 545), in the second of his three important and refreshingly sane 
papers relating to the development of skull-bones, comments upon these supposed 
fusions along the lateral line series and suggests that " fusion is a result of dermal 
bones meeting at a time when their growth-rates exceed that of the surrounding 
tissues by a certain amount. The earlier appearance of neuromast rudiments would 
increase their chances of growing to contact other neuromast rudiments and this 
would increase at least the possibihty of fusion ". Parrington goes on to quote the 
possible correlation between the increasing tendency of lateral-line elements to 
" fuse" with decrease in length of skuU, but notes the outstanding exception, in 
reverse, of the long headed Fleurantia, where " fusion " is still marked. Later he 
emphasized the possibihty of "almost complete capture of vacated territory " (1956) 
406). 

In the author's opinion Parrington's most important contributions to the questions 
concerning the growth and form of bones and their relationships to the sensory 
canals are these (Parrington 1949 : 69, 76 ; 1956 : 405, 408 etc.), which may be called 
" Parrington's axioms ". 

^ Yi and Yj are reversed from WestoU's notation so as to be in logical sequence, and I is labelled I, for 
reasons stated on p. 26. 



THE HEAD OF DIPTERUS V A LE N C I E N N E S I 13 

T. "... it is unwise to assume bone fusions without the evidence of multiple 
centres of radiation of either the ornament or the bone structure ". 

2. "... precursors of the dermal bones attract the growing primordia of the 
lateral line system and that changes in the number and positions of the dermal bones 
can therefore cause changes in the relations of the lateral line canals ". 

Examination of the partly dissected heads here figured does seem to show that if 
these two principles are accepted a reasonable interpretation can be made of the bones 
and canals, and further for this purpose Forster-Cooper's alphabetical system of 
identification of the bones, as now modified, is infinitely preferable to attempts to 
name the bones, especially as it will be shown that the fusions postulated are only too 
frequently not substantiated. 

The Sensory Canals and Pit-lines 

Before proceeding with the question of invasion or fusion of the dermal roof-bones 
it is necessary to confirm the course of the sensory canals of the pit-lines. 

The pit-lines do vary, especially the middle and posterior lines in old specimens, but 
it is a variation in detail and seldom approaches the more aberrant patterns shown by 
Forster-Cooper's (1937 : 233, text-fig. 5). The anterior line runs forwards diagonally 
from B to J, and the two sides may meet in the middle or have small loops. 

The middle line runs transversely from 1 2 into Yg and the posterior backwards 
from I2 to Z. They may be in contact in Ig and have terminal twists and curls, 
rarely partial duplication. A well marked additional line is shown in Text-fig. 42 
(P.17410). 

The general distribution of the sensory canals, as already noted, is well known, but 
not in any detail, especially in the snout region. The course of the supraorbital 
with many of the canahculi is shown in no. 33165 (Text-fig. 14), in which the canal of 
the left side has been fully developed. 

The supraorbital canal runs normally from the end of the anterior pit-hne in J to 
K, and so forwards through the usual series labelled Lj, Lj and M to N where it turns 
outwards to P. This plate is often incompletely separated in front from the snout. 
Thence the canal continues forwards, curving inwards and then again forwards and 
so downwards to the opening on the upper lip. This opening is so large in some 
specimens of Rhinodipterus (see White 1962 : 4, pi. 2, text-fig. 3) that Gross (1956 : 23, 
pi. 5, fig. 5, text-fig. 14C) identified it as the anterior nasal opening, and 0rvig (1961 : 
15. pl- 3. fig- 3. text-figs. SB, C,gC-E) as " probably openings for glands of some sort ". 

The canals of the extreme rostral region are still better demonstrated in Text-fig. 
15 (P. 6087), in which the rostral commissure has been naturally weathered out, in 
P. 34547 and in a broken section in 33166 (Text-fig. 16). In another weathered but 
much flattened snout (P. 34546, Text-fig. 17) the supraorbital canal, as it turns 
inwards in P, apparently gives off forwards six or seven pouch-like cavities instead 
of the usual rounded tubuli, and such " pouches " are also indicated in P. 34558. 
This is almost certainly the effect of crushing and weathering, accentuated by 
resorption of the bone in the immediate neighbourhood of the tubuli. An inter- 
mediate stage is shown in Text-fig. 19 (P. 46690), which also shows the presence of 
smaller openings on the hinder side of the canal. 



THE HEAD OF DIPTERUS V A LE N C I EN N E S I 




Figs. 14-17- Dipterus valenciennesi S. & M. Fig. 14, Skull-roof in which the sensory 
canals have been wholly dissected out on the left side, partly on the right. Thurso, 
Caithness. 33165. Xi-3. mil, main lateral line. For other lettering see Fig. 15. 
Fig. 15, Skull-roof in which the sensory canals of the snout have been naturally weathered 
out and some of the remainder dissected out. "Caithness". P. 6087. X2. ifc, 
infraorbital canal, roc' , rostral commissure, soc , anterior aperture of the supraorbital 
canal. Fig. 16, Median part of upper lip showing both external apertures of the supra- 
orbital canal [soc'), with part of rostral commissure [roc') and supraorbital canal itself 
and branches leading to external pores [ep) exposed by a natural longitudinal fracture. 
Thurso, Caithness. 33166. X2. Fig. 17, Skull-roof with part of canals on snout 
naturally weathered out. Foreshore, Clardon Haven, Caithness. P. 34546. (Colld. 
D. L. Dineley, 1957). ^- 1'3- '/c, infraorbital canal, soc, supraorbital canal. 



THE HEAD OF DIPTERUS V A LE N C I E N N E S I 15 

The main canal runs through Z, in which it gives off the occipital commissure, first 
to the hinder end of 1 2 and then through A. From Z the main canal normally passes 
through Y2 and Y^, to X, in which it turns down to pass through circumorbital 4 to 
form the infraorbital canal, at the same time sending a short branch to K to anastomose 
with the supraorbital canal. 

The infraorbital canal runs through the circumorbital plates 5, 6, 7, 1^, I^ up on to 
the snout, crossing the antero-lateral corners, as may be seen in most hard-snouted 
specimens. It may finish in a pore under the snout like the supraorbital canal, but if 
so the pore is often very small, and not always to be distinguished easily from the 
others present in this region; for the whole canal under pressure tends very readily to 
dissolve in the system of pores and spaces which permeate the bone there. 

To give a general impression of the canal-system, the details from Text-figs. 14, 15 
etc. have been superimposed on the left side of the outline of a well known, more 
complete skull of " platycephalus " type (P. 7834, Text-fig. 18) to which plates 
Z-A-Z have been added and, in fig. A (P. 6507), the upper lip of another specimen. 
The skull-roof has been figured by three other authors (Jarvik 1950, text-fig. 6; Gross 
1956, text-fig. 135 ; 0rvig 1961, text-fig. yB), each giving a somewhat different 
interpretation of the plates, and all different from that given here. The right side 
shows the actual distribution of the pores in this specimen. 

From this general pattern there are a number of significant deviations, as there 
are from the standard patterns of the head-plates, and since they are interdependent, 
sensory canals and head-plates wiU be considered together, in the light of Parrington's 
axioms (p. 11 supra) and of his theory of " Patterns of dermal bones " (1956). 

It is in the lateral line series, Z-X and K-P that most difficulties arise, but there 
are four plates and areas that have sufficiently constant features to make them 
readily recognizable, Z and Y^ at the back of the skull-roof and N and P near the 
front. 

Z, formerly H, is the hindmost plate and always roughly triangular. It receives 
the main-line canal from the supracleithrum and passes it on to Y2 after the canal 
has been joined by the occipital commissure. Only once in the series available is it 
in any way abnormal. 

Y^ always occupies the same position, just overlapping the front of the operculum, 
and so having a characteristic projecting angulation on the outer side. On the mesial 
side it lies alongside the hinder part of J and a small part of I2. Very rarely is it 
replaced by, or fused with, X (Text-fig. 25 ; WestoU 1949, text-fig. zB). 

N is the plate, often fused with the snout, where the supraorbital canal turns 
outwards at the start of the sigmoid curve in front, and P (WestoU's " Q ") is the 
plate or area in which the canal turns forward again. 

A survey of the available material will make clear the advantage of a purely 
alphabetical system of bone nomenclature. 

The first specimen to be considered must obviously be Forster-Cooper's standard 
specimen (P. 22187, Text-figs. 11-13). Fig. 12 shows the actual specimen as it now 
is, partly dissected. It also shows the original lettering except that G and M are 
omitted, G because it was only a scale and M because its supposed presence was due 
to a crack in L. 



i6 



THE HEAD OF DIPTERUS V A L E N C I E N N E S I 



The sensory canals are perfectly orthodox, as may be seen in the revised diagram 
(Text-fig. 13), but the series of lateral line bones show two interesting deviations, 
apart from the absence of M. A single bone occupies the space not only of M but of 
both the " L " bones and there is clearly only a single centre of growth, which from 
its central position is judged to be that of the middle bone of the three, L^. There is 




Fig. 18. Diptenis valenciennesi S. & M. Skull-roof of large head with plates Z-A-Z 
added, showing external pores of sensory canals on right side, and restoration of the 
canals on the left side, based on Figs. 14, 15. [cf. Jarvik 1950, text-fig. 6 ; Gross 1956, 
text-fig. 13B ; 0rvig 1961, text-fig. 7B). Coast near Thurso, Caithness. P. 7834. 
XI -5. A, underside of snout of similar specimen. Toldale Quarry, between Thurso 
and Reay, Caithness. P. 6507. Xi-5. ifc, infraorbital canal, mil, main lateral line. 
soc, supraorbital canal and aperture. 



THE HEAD OF DIPTERUS V A LE N C I E N N E S I 17 

also only a single Y bone, but this is judged to be compound, Y^ + Y2, by reason of 
its shape and the distribution of the pores which seem to indicate two centres. 

In 33165 (Text-fig. 14) the two sides make a most interesting contrast. On the 
wholly dissected left side a large single bone in the M-Lj area does seem to have 
had two centres and is accordingly labelled as a compound bone, and the large 
odd-shaped bone between L^ and Y^, is almost certainly a compound K -|- X. On 
the partly dissected right side the L^ to M region is similar to the left, but the K and 
X area is very different, for K is not only small but is excluded from contact with C 
by an extension of L^ and has lost to X the beginning of the supraorbital canal where 
it passes out of J. This is most unusual, but the result can be simply explained a la 
Parrington, by delay in the development of K's rudiment, which has allowed the 
capture of the beginning of the supraorbital canal by X and the invasion of this 
territory on the other side by L^. 

P. 6087 (Text-fig. 15) shows the suppression of M on both sides, in favour of N on 
the right, and probably of L 2 on the left. 

On both sides the important K-X region is occupied by a single, not very large 
bone, with a single growth centre but containing both the turn-down of the main 
canal at the beginning of the infraorbital canal, typical of X, and the continuation of 
the supraorbital canal from J, typical of K. It is not clear which has been suppressed, 
but it is probably K, for L^ on both sides is much larger than usual and extends 
further backwards into the K area to meet J, as it also does on the right side in the 
previous figure. 

In a short -headed specimen (P. 34546, Text -fig. 17) two bones only occupy the 
M-L position, each with a single centre and those of the left side are only superficially 
separated. It is probable from their forward position that the anterior bone on each 
side represents M, but that the hinder bone on the left side, judging by the position 
of the growth-centre, seems to be Lj, whereas on the right L^ is the more probable 
survivor, particularly as M is unusually large, as if it had partly replaced h^. 

The fine narrow head from near Thurso (P. 7834, Text-fig. 18, p. 13 supra) is 
undissected, the sensory canals on the left side being diagrammatic. It is one of the 
longest heads examined, 6-9 cm. from the hinder margin of B to the tip of the snout, 
being only two millimetres or so shorter than the Toldale head (P. 6507, Forster- 
Cooper 1937, pi. 7, fig. 14) and a specimen from Edinburgh (1859.33.622), but 8 mm. 
less than the record head in that Museum (1901 . 153 . 2) which measures 77 cm. The 
head in Text-fig. 18 shows quite markedly contradictory tendencies, a tendency to 
the decrease in size of plates in the middle lateral regions (K, L^, Lj) especially on the 
left side ; but in front there is a very obvious increase in size through the suppression 
of M, to the advantage of N only on the left side, but also for the benefit of Lg on the 
right. A marked difference in development of the sensory canal plates of the two 
sides is illustrated by a Clardon Haven specimen (P. 46690, Text-fig. 19). 

Of considerable interest in showing the relationship between the distribution of 
pores and the actual course of the canals is the specimen (P. 22189) on which WestoU 
(Graham-Smith & Westoll 1937 : 244, text-fig. 2a ; Westoll 1943 : 89, text-fig. yc ; 
1949 : 126, text -fig. i) based his modified scheme of lettering. 

In the original somewhat diagrammatic and restored drawings X and K are shown 
as a compound plate. The supraorbital canal entered this plate from J on its pos- 



i8 



THE HEAD OF DIPTERU S V A LE N C I EN N E S I 



terior-inner side, and almost opposite this the main canal is supposed to have come in 
from the anterior Y plate and then turn down into circumorbital " 4 " to form the 
infraorbital canal. In fact dissection (Text-fig. 20) shows that the main canal never 
reaches the " X and K " bone but turns down in Y^ (Yg of West oil), and that the 
connection between the main and supraorbital canals is not only drawn out but 
apparently double. Examination of the structure of this supposedly compound 
plate shows clearly enough that it had a single growth-centre ; X has failed to 




Fig. 19. Dipterus valenciennesi S. & M. Skull-roof showing asymmetrical development 
of lateral line plates. Clardon Haven, Caithness. P. 46690 (Colld. R. V. Collier, 1964). X2-25. 



THE HEAD OF DIPTERUS V A LE N C I E N N E S I 



19 



develop and its space has been largely taken by K, but the lateral line has been 
largely captured by Y^. The L plate is also shown as a compound 1^^ + Lg, but 
this has a single centre and seems to be composed of L^ only which has taken over the 
territory of a missing Lj. 

The pores in the very small head illustrated in Text-fig. 41 (P. 22194) seem to 
indicate a similar situation in regard to the XK area, and here also the area of M 
has apparently been taken over by Lj. 

The capture of the infraorbital curve by Y^, is e^'en more clearly demonstrated in 
Text-fig. 21 in a very abnormal short-headed fossil from Achanarras (P. 17642, cf. 




Fig. 20. Dipterus valenciennesi S. & M. Left side of skull-roof and cheek of original of 
Westoll's standard specimen (see Graham-Smith & Westoll 1937 • 244, text-fig. la ; 
Westoll 1949 : 126, text-figs. lA, C) with sensory canals partly dissected out and plates 
partly re-lettered. Achanarras, Caithness. P. 22189. X3-5. 



20 THE HEAD OF DIPTERUS V A LE N C I E N N E S I 

Forster-Cooper 1937, pi. 5, fig. 9). Plate X apparently did not develop, to j udge from 
single-centred K, but there seems to have been some opposition by K to the annexation 
of the main canal by Y^, judging by its subdivision. This fragmentation of canals 
owing to disturbance by fusion or elimination of plates is probably not unusual 
(Text-figs. 22-24). I^ 42403 (Text-fig. 22), a much resorbed specimen from near 
Wick, K and X do seem to have fused to show what appear to be double centres of 
growth, but nevertheless the start of the infraorbital canal has gone to Y^. 

In Text-fig. 21 another most interesting abnormality is illustrated. The supra- 
occipital commissure shows much disturbance owing to the non-appearance of Yg, 
but instead of its space being occupied by Y^, that bone is of the average size, and Z 
has grown forwards taking with it the outer end of the commissure, which crosses 1 2 
much further forward than usual. In contrast, the occipital cross-commissure in 
two other fishes from Achanarras (P. 17641, P. 17643, Text-figs. 23, 24), instead of its 
usual path Z-I2-A-I2-Z, misses 1 2 altogether, although these plates seem normally 
developed. 

In P . 17641 (Text-fig. 23) K and X seem undoubtedly to have fused on the left side, 
but on the right K has been eliminated to the advantage of L^, which is rather unusual 
(but see P. 6087, Text-fig. 15). Generally K is a better " stayer " than X, and it is 
worth noting that its division from J is often only superficial, the basal layers being 
completely fused (cf. Text-figs. 15, 22 etc.). On both sides the start of the infra- 
orbital canal has been largely captured by Y^ and only a small subdivision of it is 
retained by X. 

There is some doubt whether the missing plate on the right side is K or X, but the 
arrangement is partly paralleled in Text-fig. 15 where there was similar expansion of 
Li into the K area and disarrangement of the canals. L^ and L2 in P. 17643 (Text- 
fig. 24) show a very obvious case of imperfect fusion. 

Of the two " brachypygopterus " types from Banniskirk illustrated (42480, 
R.S.M. 1859.33.624, Text-figs. 5, 6), the first shows a fairly obvious X and K fusion 
and the L^-Lg, M and possibty N area is occupied by a single plate. The second 
shows a large N and on the left side only one small plate (labelled L^) between N and 
K, also a large Y^ and no Yg. The naturally exposed sensory canals are standard as 
far as they may be traced. 

Attempts to Identify the Head-plates 

The difficulty of tracing the canals by the pores led Stensio {in Holmgren & Stensio 
1936 : 366, text-fig. 280^-C) to misplace the connection between the supraorbital 
canal and the main canal in the three rather difficult Edinburgh skulls on which he 
based his nomenclature (Text-figs. 25-27). 

Stensio's choice of a key-specimen (R.S.M. 1878. 5. 164) on which to base his 
identifications was not altogether fortunate, for it suffers very much from multiphca- 
tion of plates in the central area and in the front lateral line series. His figure (Text- 
fig. 28), was formalized and partly restored, cf. Text-figs. 25, 29. 

Neither of Stensio's supraorbital bones (SO^, SO2) belongs to the circumorbital 
series, but his " dermosphenotic + postorbital " {Dsph + Po) does, and the orbit 
is farther out and certainly smaller. This bone is not compound but is the circum- 
orbital 3 of Forster-Cooper's notation. In the original specimen it is preserved only 



THE HEAD OF DIPTERUS V A LEN C I EN N E S I 




Figs. 21-24. Dipterus valenciennesi S. & M. Fig. 21, Left posterior part of skull-roof and 
cheek with sensory canals dissected out (cf . Forster-Cooper 1937, pi. 5, fig. 9) . Achanarras, 
Caithness. P. 17642. X<4. CI, cleithrum. Op, operculum. Orb, orbit. Sc, scale 
(Forster-Cooper's "G"). SCI, supracleithrum. s.com, occipital cross-commissure. 
Fig. 22, Posterior half of much resorbed skull-roof with sensory canals partly dissected 
out to show probable fusion of plates K and X on each side. Killimster, near Wick, 
Caithness. 42403. X4. Fig. 23, Skull-roof with sensory canals dissected out (cf. 
Forster-Cooper 1937, P^- 6, fig. 11 ; Westoll 1949, text-fig. 2E). Achanarras, Caithness. 
P.17641. X2-5. s.com, occipital cross-commissure. Fig. 24, Right posterior part of 
skull-roof with sensory canals dissected out (cf. Forster-Cooper 1937, pi. 4, fig. 8). 
Achanarras, Caithness. P. 17643. X2. ifc, infraorbital canal. LLSc, lateral line 
scale. Sc, body scale. SCI, supracleithrum. s.com, occipital cross-commissure, 



THE HEAD OF DTPTERUS V A LE N C I E N N E S I 




Figs. 25-27. Diptenis valenciennesi S. & M. Camera lucida drawings of originals of skull- 
roofs of " platycephalus " type, sketched by Stensio (in Holmgren & Stensio 1936 ; 366, 
text-figs. 280A-C). Fig. 25, Large head (6-i cm. long) with median mosaic and unusually 
numerous plates (6) between K and P. Locality unknown. R.S.M. 1878.5. 164. 
X I -3. Fig. 26, Medium sized head (4-6 cm. long) with anterior mosaic and only three 
plates between K and P on right side. Firth, Orkney. R.S.M. 1898. 163.6. xi-3. 
Fig. 27, Large head (6-3 cm. long) showing tendency to subdivision of plates between K 
and P. Locality unknown. R.S.M. 1859.33.32. X i -3. 



THE HEAD OF DTPTERUS V A LEN C I EN N E S I 23 

on the left side and carries an unusually large branch of the supraorbital canal, for as 
a rule the pores are few, often absent. This skull-roof, in spite of its oddities, is 
reasonably reduced to order under Parringtonian influence with Forster-Cooper's 
amended lettering (Text-fig. 25). The only point of real importance is that Y^, 
normal on the left side, has most exceptionally not developed at all on the right side, 
where its place has been covered very largely by a much expanded X, and only to a 
small degree by the advance of Yg. 

Both of Stensio's other two specimens show peculiar features. The small broad 
head (R.S.M. 1898. 163. 6, Text-fig. 26) shows division of J on the left, and no M and 
a possible fusion of K and X on the right ; the larger head (R.S.M. 1859. 33. 32, 
Text-fig. 27) shows fusion of K and X on the left side, fusion of L^ and Lg and im- 
perfect subdivision of N on the right. 

Lehman (1959) in his work on the Upper Devonian Dipnoans from Greenland 
followed Stensio in endeavouring to name the plates of the skull-roof, although by no 
means agreeing on the terms to be used, nor for that matter on the homologies when 
he did. 

Comparison of Text-figs. 28-30 well illustrates the confusion that can be caused by 
efforts to identify these head-plates with the bones of other fishes. 

Stensio's supposedly compound " dermosphenotic and postorbital " [Dsph + Po, 
Text-fig. 28) is not the homologue even in part of Lehman's " Dermosphenotique " 
[Dsph, Text-fig. 30), as a glance at Text-fig. 29 will show, for Stensio's plate is a simple 
circumorbital, no. 3 of Forster-Cooper. Lehman's " Dermosphenotique " is a single- 
centred plate (Lehman 1959, pis. 1-14, 16) and represents the survivor of K or X, 
probably K, which has taken over the other's territory and acquired all the canals 
(of. Text-figs. 19, 20, 24). It is therefore strictly homologous with the most posterior 
member of Stensio's " laterale Frontalserie " {Fr. I, Text-fig. 28) of which Lehman's 
" Nasaux " are the front members. 

Again, Stensio's supposedly compound " Supratemporo-Intertemporale " (It -\- 
St, Text-fig. 28) is also a simple bone, the X of Text-figs. 25, 29, which has invaded 
Yj's territory, and Y^, the equivalent of Lehman's " Dermopterotique anterieur " 
[Dpt. i) has most exceptionally not developed at all. But Lehman's " Dermo- 
pterotique posterieur" [Dpt. 2) is the Y2 of Text-fig. 29, and thus the homologue of 
Stensio's " laterale freie Extraskapularplatte " {Ext. 1 2, Text-fig. 28). This last 
plate, which is laterally imperfect in the original (Text-fig. 25) does not show any 
part of the supratemporal cross-commissure as indicated by Stensio (Text-fig. 28) ; 
it would have been present on the plate behind (Z), which is missing. 

Stensio's " hintere laterale Parietalplatte + einer lateralen Extraskapularplatte " 
{Pa. I2 + Ext. Ij^, Text-fig. 28) which is identical with Lehman's " parietolatero- 
extrascapulaire " {PalExSc, Text-fig. 30 ; Lehman, 1959 : 18, pis. 1-21, text-figs. 
2, 4, 5, 7-13, 21-23, 26), presents a very different problem, and indeed provides the 
key to the homologies of the plates of the earlier Dipnoi. These paired plates are 
the 1 2 of Text-fig. 29 and show in the available material of Dipterus and Lehman's 
illustrations a single growth centre ; nevertheless the suggestion that there is an 
extrascapular element is supported by Westoll (1947 : 134, text-fig. 3/) who states 
that " it is likely that at least bone I also arose from more than one rudiment since a 
few specimens show ossicles enclosing the appropriate section of the canal ", And 



24 



THE HEAD OF DIPTERUS V A L E N C I E N N E S I 




Fig. 28. Dipterus valenciennesi S. & M. Right side of Stensio's diagram, text-fig. 280A 
(see Fig. 25), with original lettering. 

Dsph + Po, Dermosphenotiko-Postorbitale ; Ext. h, laterale freie Extraskapular- 
platte ; Fr.c, zentrale Frontalserie ; Fr.l, laterale Frontalserie ; Fr.m-^, Fr.m^, Flatten 
der medialen Frontalserie ; It, Intertemporale ; It + St, Supratemporo-Intertemporale ; 
Pax, zentrales Parietale ; Pa.l^, vordere Platte der lateralen Parietalserie ; Pa.l^ + 
Ext.li, laterale Parietalplatte + einer lateralen Extraskapulai-platte ; Ptr.^, hintere 
Postrostralia ; SOj, SOj, Flatten der Supraorbitalserie (S.O. ( = 3) is omitted in Fig. 30}; 
ap, vordere " Pitlinie " ; ifc, Infraorbitalkanal ; mp, mittlere " Pitlinie " ; pp, hintere 
" Pitlinie " ; s.com. Supratemporalkommissur ; soc. Supraorbitalkanal. 



THE HEAD OF DIPTERUS V A LEN C I EN N E S I 



25 




r<: 30 



Plr 




DSph 



v-Dptj. 



'^PalE^Sc 



Pac 



Fig. 29. Dipterus valenciennesi S. & M. Right side of same diagram (Fig. 28) partly 
redrawn from original specimen (cf. Fig. 25), with new interpretation of sensory canals 
and plates re-lettered according to modified FCAN. Other lettering as in Fig. 28. 

Fig. 30. Soederberghia groenlandica Lehman. Right side of diagram of skull-roof by 
Lehman (1959, text-fig. 2) showing original lettering (with broken indication lines) and 
also modified FCAN. 

Dpt^, dermopt^rotique ant6rieur ; Dpt^, dermopt^rotique post6rieur ; DSph, dermos- 
ph^notique ; Frb, composant antdrieur distinct du frontal lateral ; Frl, frontal 
lateral ; Frm, frontal m6dian ; Naj-lSSsig, nasaux ; Orb, orbite ; Pac, parietal central ; 
Pal, parietal lateral ; PalExSc, parieto-lateroextrascapulaire ; Pi, plaque pin^ale ; Po, 
postorbitaux ; Ptr, postrostral ; So, supraorbital. Sensory canals as in Fig. 28. 



26 THE HEAD OF DTPTERUS V A LEN C I EN N E S I 

indeed the Belgian specimen of Rhinodipterus secans (White 1962 : 3, pi. i, text-fig. i) 
shows a similar " extra " plate enclosing the canal. Subdivision or proliferation of 
lateral line and general plates is not necessarily significant except that perhaps it 
points to the ancestral mosaic, and is of common occurrence in Diptenis, as the front 
of Stensio's specimen shows. But in this instance it throws light on the presence of 
part of the commissure on 1 2 which is anomalous, for it is clearly out of line. This 
points to the former presence of a plate on each side between A and Z, now lost, and 
that the section of the cross-commissure belonging to it has been captured by 1 2, 
that is, usually, for as noted above (p. 18, Text-figs. 23, 24) the commissure does 
sometimes go straight across through Z-A-Z, as it must have done when there were 
five plates in the " extrascapular " series, even as Save-Soderbergh (1932 : 98, text- 
figs. 18, 20) postulated for his " common ancestor " of the ichthyostegids and crossop- 
terygians ; but it was not the bone 1 2 (Save-Soderbergh's and Stensio's Pa.l^ + 
Ext.l-yj that belonged to that row and originally bore the commissure, but a plate 
behind it, represented rarely by the little plate I^, noted by Westoll, and seen also 
in Rhinodipterus. 

The above correlations may be summarized as follows : 



Stensio 1936 


Forster-Cooper 1937 emend. 


Lehman 1959 


(Text-fig. 28) 


(Text-fig. 29) 


(Text-fig. 30) 


Dsph -H Po 


(CO) 3 


SO (omitted) 


[Posterior) Fr. I 


K 


DSph 


It + St 


X 


absent 


absent 


absent (YJ 


Dpt, 


Ext. l^ 


Y. 


Dpt, 


Pa.l^ + Ext. li 


I2 


PalExSc 



To trace back the homologues of the skull-plates of Dipterus in the earher genera 
of dipnoans, the two species of Dipnorhynchus from the Lower Devonian (Siegenian)^, 
presents considerable difficulties. The roof-pattems of D. siissmilchi and D. lehmanni 
are reasonably well known, thanks to E. S. Hills (1941, pi. 9, text-fig. 5 ; 1943, 
text-fig. iB), Westoll (1949 : 140-143, text-fig. 5) and Lehmann & Westoll (1952, 
pi. 24, fig. A, text-figs. 4^, 5^). 

The most obvious points about the pattern of these skull-roofs (Text-figs. 31, 34) is 
the presence of a pineal foramen and the separateness of the supraorbital and infra- 
orbital sensory canals. The back row of plates, presumably the extrascapulars, 
were finally shown in D. lehmanni (Text-fig. 31) to consist of two pairs of plates 
identified as H(Z), the smaller outside pair, and I, the large median pair, instead of 
the Z-A-Z arrangement of Diptenis. The occipital cross-commissure, apparently 
not seen in either specimen is assumed to run across H(Z)-I-I-(Z)H. There is no 
A plate but that immediately in front, a smaUish median element, is considered to 
be B, and in front of that again a pair of larger plates completely separate in the 
German species, but partly attached to one another in the Australian. Westoll 
(1949 : 141) has resisted the temptation to label these as the " C " pair, as at first 
sight would seem obvious, on the grounds of their posterior relationship to the orbits 
and then to bones of the X-K area. This pair is labelled Bg and along side the B 

1 G. M. PhiUp & A. E. H. Pedder (1964 : 1323). 



THE HEAD OF DIPTERUS V A LEN C I EN N E S T 



27 



plates and bearing the supraobrital canal are L, K and three J plates on each side, 
separate in D. siissmilchi, or partly fused in D. lehmanni, the most posterior of the 
J plates being very large (Lehmann 1956, text-figs, i, 2). It is unexpected that B 
should be in three parts, two paired, and at least two other arrangements seem 
possible. 

Taking first the obvious one rejected by WestoU (Text-fig. 32) one might consider 
that in the later Dipterus the orbits had moved backwards from their position in 
Dipnorhynchus , bringing them opposite the anterior ends of the C plates (Text-fig. 
35), and this would account for the difference in the relative position of K, X etc. 
in the two genera; and their movements outside of J might have been a factor 




Figs. 31-33. Dipnorhynchus lehmanni Westoll. Fig. 31, Reconstruction of skull-roof 
after Lehmann & Westoll (1952 : 411, text-fig. 5A) with the original lettering (anterior 
Y-plates should be Y2, posterior, Yj). Fig. 32, The same, but re-lettered according to 
first alternative. Fig. 33, the same, but re-lettered according to second alternative. 



28 



THE HEAD OF DIPTERUS V A LEN C I E N N E S I 





Fig. 34. Dipnorhynchus sussmilchi (Etheridge). Outline reconstruction of skull-roof, 
after HQls (1933 : 637, text-fig. 2) with addition of the sensory canals from Hills (1941 : 
646, text-fig. i) and the back of the skull-roof restored in outline, after Westoll (1949 : 
142). The lettering according to second alternative (cf. Fig. 33). 

Fig. 35. Dipterus valenciennesi S. & M. Reconstruction of skull-roof, after Graham- 
Smith & Westoll (1937, text-fig. aa) and Westoll (1949, text-fig. lA), the lettering modi- 
fied after Fig. i8. 

Fig. 36. Scaumenacia curta (Whiteaves). Reconstruction of skull-roof, after Westoll 
(1949, text-fig. 6D) with the lettering modified. 



THE HEAD OF DIPTERUS V A LE N C I EN N E S I 



29 




38 



^ t^ 






Fig. 37. Ctenodus cristatus Agassiz. Reconstruction of skull-roof, after Westoll (1949, 
text-iig. B) with the lettering modified, and Watson & Gill (1923, text-fig. 21). 

Fig. 38. Ceratodus sturi Teller. Reconstruction of skull-roof, after Westoll (1949, text-fig. 

9A), with lettering modified. 

Fig. 39. Ceratodus formosus Wade. Reconstruction of skull-roof, als, approximate 

anterior limit of scale covering. 

Fig. 40. Neoceratodus forsteri (Krefft). Skull-roof, after Holmgren & Stensio (1936, text- 
fig. 288B), re-lettered. 



30 THE HEAD OF DIPTERUS V A LEN CI EN N E S I 

contributory to the anastomosing of the infraorbital and supraorbital canals. The 
paired " Bg " plates would then be C plates and instead of a proliferation of J plates 
we would have a more normal series of K, J and I. B would be small and A un- 
accounted for, unless it was the result of fusion of the very large paired plates labelled 
I by Westoll, for it seems unlikely in any event that the supposed I plates would later 
move forwards from the " extrascapular " row to embed themselves in the skull-roof 
between B and Y as in Dipterus. 

On the other hand, if we accept the possibility that the two anterior Bg plates of 
WestoU's interpretation did fuse to form the B of Dipterus, as seems indicated in 
Dipnorhynchus siissmilchi (Text-fig. 34), then an even more interesting situation 
could have held (Text-fig. 33). B, of Westoll, would be A and Jg would be Ig 
(missing on the right side) with J3 as I^, the plate that is still occasionally preserved 
as a small entity carrying part of the occipital cross-commissure in Rhinodipterus 
(White 1962, pi. I, text-figs, i, 2) and Dipterus (Westoll 1949, text-fig. 3/). On the 
outside of it is Z(H). 

At this level of development, which Westoll (1942 : 142) claims is a fair approxima- 
tion to the hypothetical ancestral condition, this row did not carry the cross- 
commissure, which was supposedly borne by the row behind, H-I-I-H of WestoU 
(Text-fig. 31), but here supposed to be a row of plates (/5-a-a-y^, Text-figs. 33, 34) 
that has now disappeared in other known fishes. This is in keeping with the 
important point shown by Westoll (1938, text-fig. 2) in his brief paper on the ancestry 
of the Tetrapods, that in vertebrates there is a movement backwards of the bones of 
the skull roof with additions at the anterior end by orderly development from the 
variables, and a loss of those behind with a shortening of the occipital region, features 
shown in some degree by Dipterus relatively to Dipnorhynchus. 

Between Dipnorhynchus and Dipterus there must have been a stage when the 
hindmost row fi-a.-a.-P was lost and the cross-commissure had been taken over by 
the full " extrascapular " row Z-I^-A-Ii-Z, before the two I^ plates lost their 
territory to their neighbours and their remnants fused with lo in front taking with 
them a segment of the commissure — as noted above, only rarely does this remnant 
of Ii ever show itself as a minute independent plate. 

From Dipterus one can move forward in time through a perfect morphological 
series based on WestoU's (1949, text-figs. 6Z), 85, 97!) outhne restorations of the 
later dipnoans Scaunienacia, Ctenodus and Ceratodus, which show progressive 
diminution and final loss of the " extrascapular " Z-A-Z row, with the cross- 
commissure being finally captured by Y^-B-Yi (Text-figs. 35-39). At the same time 
the series shows one other powerful trend coupled with the relative movement back 
of the head plates, and that is on simplification of the pattern, by invasion and loss 
rather than by fusion as Westoll indicates by his lettering. This trend is after all a 
continuation of the process seen in passing from Dipnorhynchus to Dipterus. The 
apparently unstable many plated mosaic inherited from the ancestral form persists 
in the CDE area in Dipnorhynchus ; in Dipterus the C area generally, and that of D 
and E quite often, has settled down in a more constant pattern of a few large plates, 
which is characteristic of the later terms in the series. It is interesting to note that 
in C. formosus Wade (1935 : i, pi. i ; see also Text-fig. 39) the dermal skull pattern 
can be reasonably interpreted in a form comparable with that of C. sturi Teller from 



THE HEAD OF DIPTERUS V A LE N C I EN N E S I 



31 



slightly earlier beds in the Austrian Alps (Text-fig. 38) , the most striking difference 
being that F has increased in size at the expense of D, which in the living Neoceratodus 
has disappeared altogether (Text -fig. 40). 
The various trends in this morphological series may be summarized as follows : 







Posterior 


Centre 


Lateral line 






row of 


plates 


plates, 




Age 


plates 


front to rear 


/3-X, a-P 


Dipnorhynchus lehmanni 


late 

Lower 

Devonian 


a^fia 


Ant. mosaic 
BAa 


4. 4 


D. sussmilchi 


do. 


a/3/Ja 


Ant. mosaic 
BAa 


3. 7 + 


Dipterus 


Upper 
M.O.R.S. 


ZAZ 


Ant. mosaic 

CBAor 
FEDCBA 


4. 7 


Scanmenacia . 


Early 
U.O.R.S. 


ZI2AI2Z 


ECBA 


4. 3 


Cienodus 


U. Carbo- 
niferous 


Y2ZI2BI2ZY2 


FEDCB 


4. 3 


Cerafodus sturi 


M. Trias 


YiBYi 


FDB 


I. 3 


C.formosus . 


M. Trias 


YiBYi 


FDB 


I. 3 


Neoceratodus . 


Living 




FB 


ii I 



The Endocraniunt 

The undersurface of the skull of Dipterus has been tolerably well known since the 
days of Hugh Miller (1849 : 62, text-fig. 20). Pander (1858 : 10, pi. 3, figs. 11, 13, 14) 
was the first to deal seriously with this part ; Traquair (1878 : 5, pi. 3, fig. i) 
established that the suspension was autostylic ; then Woodward (1891 : 234, text- 
fig. 36(1)) gave a restoration of the upper and lower jaws, and Goodrich (1909 : 242, 
text-fig. 2io.<4) one of the whole palate. Finally Watson & Day (1916 : 29, text-fig. 
6) published a general description of the palatal aspect of the head with an unlettered 
restoration which has been freely copied by later authors, either as originally 
published (Holmgren & Stensio 1936 : 365, text -fig. 279) or somewhat modified 
(Graham-Smith & WestoU 1937 : 251, text-fig. 8D). Save-Soderbergh (1952 : 22) 
noted further details ". . . similar canals [to those of Chirodipterus] for the efferent 
pseudobranchial arteries, grooves for the lateral dorsal aortae and orbital arteries, 
canals for occipital arteries, and grooves for the internal jugular veins ", but he gave 
no figures. Two years later Jarvik (1954 : 69, text-fig. 36B) figured a fine specimen 
in which he labelled the foramen for the occipital artery, a large paired fossa on the 
lower side of the otic region, and the groove for the lateral dorsal aorta. 

Kesteven (1951 : 108) has given a clear warning against the too positive identifica- 
tion of foramina and other features in fossil skulls. The force of this warning is 
proportional to the distance in the relationship between the fossil and a living species. 
In the case of the Dipnoi, it can be shown that the endocranium of Dipterus is 
sufficiently close to the only other described fossil Dipnoan skull, that of the Upper 



32 



THE HEAD OF DIPTERUS V A LEN C I E N N E S I 



Devonian Chirodipterus (Save-Soderbergh 1952)^ and to that of the hving Neoceratodus 
(Gtinther 1871 ; Huxley 1876 ; Kesteven 1931 ; Holmgren & Stensio 1936 : 372) 
that comparisons may be reasonably made. 

The most important specimen of Dipterus is an incomplete but uncrushed skull 
from "Caithness", P.17410, with a curiously asymmetrical pattern to its roof- 
plates (Text-figs. 42-47). This has lost the snout and most of the right side of the 
skull roof and endocranium, but the left side is reasonably well preserved, although 
the perichondral bone has flaked off in places and intractable matrix obscured 
others. The articular head of the left palatoquadrate is displaced upwards. This 




Fig. 41. Dipterus valenciennesi S. & M. Head of small specimen showing part of lower 
dentition of each side. Achanarras, Caithness. P. 22194. ^5- ^'^. anterior gular ; 
Br, branchiostegal ray ; CI, cleithrum ; LatL, lateral line ; Op, operculum ; PG, 
posterior gular ; SOp, suboperculum. 

' Save-Soderbergh pointed out (p. 8) that the snout of Chirodipterus if broken off would be identical 
with the fossils described as Ganorhynchus. The locality and formation of the unique holotype (44627) 
of the type-species, G. woodwardi Traquair (1873 : 555, pi. 14) are unrecorded, but Mr. H. A. Toombs 
has established that the matrix is so similar to that of Middle Devonian (Couvinian) fossils from the 
neighbourhood of Gerolstein in the Eifel as to leave little doubt as to its source. 



THE HEAD OF DIPTERUS V A LEN C I EN N E S I 



33 



skull was most probably developed by Save-Soderbergh and formed the basis of his 
brief remarks on the endocranium of Dipterns (1952 : 22) in his classical description 
of the skull of Chirodipterus. 

The bony structure of the endocranium is exactly as that described in Chirodipterus 
(Save-Soderbergh 1952 : 6) consisting of inner and outer shells of thin laminar peri- 
chondral bone which also hned the vascular and nerve canals connecting the two 
capsules. The space between is largely filled by cancellar bone. The latter is 
frequently exposed in this specimen owing to the loss of the perichondral layer, and 
it is not always a simple matter to determine whether the supposed foramina are 
genuine or not. As often observed in this genus and noted in Chirodipterus the 
neurocranium is undivided and forms a single unit with the palatoquadrates which 




Fig. 42. Dipterus valenciennesi S. & M. Imperfect and irregular skull-roof. Same specimen 

(P. 17410) as in Figs. 43-47. X2-5. 



34 



THE HEAD OF DIPTERUS V A L E N C I E N N E S I 








Figs. 43-47. Dipierus valenciennesi S. & M. Imperfect skull. Locality in Caithness 
unknown. P. 1 7410. Fig. 43, Left side view. Palatoquadrate removed. X2-5. 
Fig. 44, Planum antorbitale. x <4. Fig. 45, Outline of left orbitotemporal space. 
X2-5. Fig. 46, Three-quarters ventral view. X2-5. Fig. 47, Back of skull. X2.5. 
a.occ, foramen for occipital artery ; a.or, foramen for orbital artery ; a.or.p, posterior 
opening for orbital artery ; aps, posterior opening of efferent pseudobranchial arter^"^ ; 



THE HEAD OF DIPTERU S V A LE N C I E N N E S I 35 

form on each side a high transverse lamina, in this specimen with a forward slope of 
about 45°. A major difference between the skulls of Dipterus and Chirodipterus is 
that in the former the neurocranium is in contact with the cranial roof throughout 
its length, so there is no median fossa or crest over the otic and occipital regions, the 
fossa for the temporal muscle {tf) on each side being bounded by the neurocranial 
wall medially and by the thin dorso-lateral crest {die) on the outer side (P.17410, 
Text-fig. 47). This crest is probably continued backwards under the " tabular 
horn " which is a projection of plate 1 2 itself and seems to comprise all three layers 
(P. 46761, PI. 3, figs. I, 2). 

The lateral crest (/c), in continuation of the palatoquadrate, forms the outer wall 
of the masseter fossa (m/) on the inside and the roof of the branchial cavity on the 
outer. 

Of the ethmoid region in front of the planum antorbitale there remains only a short 
length and cross-section of the perichondral lining of the right olfactory canal {nl, 
Text-fig. 46), through the post -nasal wall, and it is clear that, as in Chirodipterus, 
ossification was in this part only perichondral. The anterior ramifications of the 
profundus in the roof of the nasal capsules along with other canals and vessels, are 
well displayed in R.S.M. 1859.33.612, and to a lesser extent in P. 46693. 

The ethmoidal part of the orbital space, preserved only on the left side, is very 
different from that of Chirodipterus or Neoceratodus (Text-figs. 43-45), for it is much 
longer, occupying as much of the space as the orbito-temporal region. The posterior 
face of the planum antorbitale is completely ossified and runs diagonally from the 
outer margin backwards and inwards to the neurocranial wall. The outermost part 
is formed by a very large buttress (b-^ which joined the entopterygoid to support the 
dental plate. Narrow at the base it had a wide triangular section dorsally and a 
small space-filling talon [ta) outside it. Separated from it by a deep cavity containing 
the main profundus foramen {prof) was an even larger multiple buttress {b^ forming 
the neurocranial wall and pierced by two or three small foramina for branches of the 
profundus. The lower half of this region, which is probably a little crushed down- 
wards, is obscured by debris {de), but a diagonal fracture in P. 755, apparently 
through the second buttress immediately above the inner posterior angle of the left 
tooth-plate shows the buttress passihg below into a horizontal lateral shelf above 
the entopterygoid. 

These buttresses provide a necessary support, from the skull-roof, to massive 

art.s, articular surface of palatoquadrate ; b^, b^, buttresses ; ch, notochord ; c.i, groove 
for internal carotid ; db, debris ; div.end, diverticulum of endolymphatic duct ; die, 
dorso-lateral crest ; dpt, tooth-plate ; Enpt, entopterygoid ; for.m, foramen magnum ; 
hm, exit of truncus hyomandibularis VII ; l.c, lateral crest ; lev, foramen for lateral 
cerebral vein; mf, masseter fossa ; Pq, palatoquadrate (cut away in Fig. 43) ; prof, 
main profundus foramen ; prof -\- vju, foramen for profundus nerve and jugular vein ; 
Psph, parasphenoid ; s.a, groove for lateral dorsal aorta ; ta, talon of buttress ; tf, 
temporal fossa ; vca, foramen for anterior cerebral vein ; Ve, fused vertebra ; vju, groove 
for internal jugular vein ; v. pit, groove and foramen for pituitary vein. 
nl, olfactory nerve; II ; III ; IV; Fj, 3, opening for mandibular and maxillary 
branches of trigeminal nerve ; VIIIp, posterior branch of acoustic nerve ; IX ; X ; 
Xbr, branches of vagus. 

3§ 



36 THE HEAD OF DIPTERUS V A LEN C I EN N E S I 

dental plates, which with their strongly denticulated surfaces, were clearly capable of 
dealing with very rough food. Mollusca are not very evident in these beds, and 
such denticulated plates usually with no obvious facets of wear, speak rather for 
gripping and crushing struggling animal prey than for cutting inert vegetable matter, 
so arthropods, worms and perhaps other fishes suggest themselves. That Chirodi- 
pterus had feebly sculptured plates and rudimentary buttresses — Save-Soderbergh 
(1952 : II, pi. 3, fig. 2) speaks only of " a vertically elongated prominent area " — 
suggests that it fed on softer unresisting matter such as carrion. On the other 
hand the earlier Australian marine Dipnorhynchus had a large free buttress at the 
outer margin of the entopterygoid in the middle of the orbito-temporal space 
(P . 33699) to support its large massive dental plates for holding and crushing shelly 
prey, such as the numerous brachiopods with which its remains are associated. 

Only the outer face of the left wall of the orbito-temporal region of the neuro- 
cranium of Dipterus is reasonably preserved (P. 17410, Text-iig. 43). The brain-case 
is short and narrow, and the wall, as preserved, curves gently inwards towards the 
skuU roof. There is a wide subocular shelf, a continuation of that already noted, and 
this passes imperceptibly into the processes of the palatoquadrate. In front high up 
and just behind the second buttress is a large foramen for the anterior cerebral vein 
{v.ca). Below and behind this is a very large dumb-bell shaped aperture for the 
optic nerve (//). Whether this is its true shape is not quite certain, but if so, it is 
unlike what is found in either Chirodipterus or Neoceratodus. As in Chirodipterus, 
above and behind in a slanting hne are three foramina, the topmost for the fourth 
nerve {IV), the middle and largest for the oculomotor (///) ; and on a level with the 
foramen for the optic nerve, a groove and foramen for the pituitary vein [v. pit). 
Immediately behind and below this vein is the large foramen for the profundus nerve 
and the internal jugular vein {prof-{-v.jn). This foramen hes at the junction of the 
endocranium and the fused processes of the palatoquadrate and faces forwards. 
High above this foramen is a large opening corresponding to that identified in 
Chirodipterus for the mandibular and maxillary branches of the trigeminal nerve 
(F2,3), but it is relatively farther back, and below it, instead of at the side, is the 
smaller foramen for the orbital artery (a.or). 

Behind the palatoquadrate on the left side the ventro-lateral surface of the otic 
and occipital regions is reasonably well preserved (Text -figs. 43, 46, 47), although the 
upper part formed by the lateral crest [Ic) is possibly a little distorted by crushing. 
Below it the wall of the endocranium is almost straight and vertical except for the 
slight swelling over the saccular division of the labyrinth cavity. 

The back of the skull slopes somewhat irregularly at an average angle of 60° to 
the base of the occipital region as preserved, but although two vertebral elements 
[Ve) are seen to be incorporated in this part it is clearly imperfect. 

A number of features are readily identifiable with those recognised in Chirodipterus. 

Immediately above the labyrinth swelling lies the groove for the internal jugular 
vein (yjn) which leads to the foramen piercing the palatoquadrate, and just in front 
of this foramen is another, almost as large, probably for the exit of the truncus 
hyomandibularis VII{hm), and below it the posterior opening of the canal for the 
orbital artery (a.or.p) to which a groove leads from that of the dorsal aorta (s.a). 



THE HEAD OF DIPTERUS V A LE N C I E N N E S I 37 

Where these two grooves meet a third groove, for the internal carotid artery, passes 
sHghtly medially into a notch {ci) and then under the parasphenoid. Above it is 
the posterior opening of the efferent pseudobranchial artery (aps) . All these features 
are little more than a paraphrase of what has been found in Chirodipteriis, but not 
seen in that genus is a very large opening, possibly enlarged by fracture, just behind 
the junction of these same three grooves, probably for a posterior branch of the 
acoustic nerve {VI Up). Above and behind this is the foramen for the glosso- 
pharyngeal nerve {IX) . 

The short occipital region is continuous with the otic but the sides are flatter and 
more nearly vertical. Owing to the slope of the back of the head, about 60° but 
exaggerated by breakage, the dorsal flange formed by the lateral crest is present only 
in the anterior part. 

At least two vertebrae are fused on to the basioccipital region, and in other 
specimens (DMSW.P.149, PL i, fig. 2; also P. 755 and R.S.M. 1859. 33. 617) it seems 
evident there are three. 

The main features of the otic region, the internal jugular groove {vju) above and 
that of the lateral dorsal aorta {s.a) below, are continued on to the occipital region. 
The vagus foramen {X) is in the jugular groove in much the same position as in 
Chirodipterus (Save-Soderbergh 1952 : 19, text-fig. 8). Immediately above and at 
right angles to it is a large foramen running into the back of the skull, presumably 
for the lateral cerebral vein {lev, Text-figs. 43, 47), and above this are two smaller 
openings for branches of the vagus {Xbr). However, the most conspicuous feature 
in the back of the skull is a pear-shaped opening in the centre near the skull-roof, 
presumably a cross-section of a median diverticulum of the endolymphatic duct, 
{div. end) as in Eusthenopteron (Stensio 1963 : 97, 100, text-figs. 50A, 51A). 

The groove of the lateral dorsal aorta {s.a) widens to the rear and joins that of the 
right side behind the parasphenoid (P. 34549, Text-fig. 49 ; cf. Jarvik 1954, text- 
fig. 36B). Just before it does so, it is pierced at its upper margin by a foramen for 
the occipital artery {a.occ), which is nearly horizontal owing to the swelling out of 
the bone. It penetrates the bone for almost 3 mm. by a somewhat sinuous course, 
and re-emerges above and slightly behind by a similar foramen above, then continues 
upwards by a well-marked groove (P . 34544 ; also DMSW . P . 149) which apparently 
bifurcates before the vessel re-enters the skull through two distinct foramina (Text- 
fig. 48). 

The right side of the occipital region of P.17410 is too imperfect to show these 
features, but the vagus foramen, much enlarged, is present and a little below and 
behind it, as in Chirodipterus, two or three foramina for spino-occipital nerves. 

Of the dermal bones of the palate in this specimen, only the right half of the 
parasphenoid and part of the right entopterygoid is preserved. The former is sunk 
into the endocranium so that the surfaces are level, and they give the impression of 
being joined by a suture ; and the entopterygoid, to which the parasphenoid is 
truly sutured, has the same appearance in regard to the palatoquadrate. 

The features of the palate are better seen in other specimens, in particular P . 34549, 
a specimen collected by Dr. D. L. Dineley in Clardon Haven, near Thurso (Text-fig. 
49, PI. I, fig. i). It is a crushed head with very well preserved, but flattened 
palate. 



38 



THE HEAD OF DIPTERUS VALENCIENNESI 



The anterior angle of the parasphenoid in this specimen shows a very well marked 
hypophysial foramen (hy.f) which runs inwards (upwards) and somewhat backwards. 
There is a broad, fiat or slightly concave selvage along the antero-lateral sides which 
is widest in front medially and decreases laterally to vanish at the greatest breadth. 
On the medial side of this selvage the parasphenoid shows a complementary depres- 
sion or groove, narrowest medially in front and widest behind at the lateral angle. 
Between these grooves the parasphenoid is gently convex. Near the front of this 




'\yff--- 



.^ 



Figs. 48, 49. Dipterus valenciennesi S. & M. Fig. 48, Basioccipital region to show course, 
partly excavated, of occipital artery {a.occ). Clardon Haven, near Thurso, Caithness. 
P. 34544. X2-25. Fig. 49, Palatal view. See also PI. i, fig. i. Clardon Haven, near 
Thurso, Caithness. P. 34549. X2-25. Both specimens colld. D. L. Dineley, 1957. 
fd, fold ; hy.f, hypophysial foramen ; s.d, secondary denticles. Other lettering as in 
Figs. 43-47. 



THE HEAD OF DIPTERUS V A LEN C I EN N E SI 39 

median area and some distance behind the hypophysial foramen there is a very 
conspicuous crescentic, transverse sHt or infolding of the surface of the parasphenoid 
directed forwards [fd] with a corresponding broad and shallow depression behind it 
in which are four or five well marked denticles on low ridges. The depth of the 
pocket cannot be seen, but probably it does not extend far in. It is seen in a few 
other specimens, in varying degrees of development ; well developed although partly 
masked by transverse cracks in P. 755, P. 34556 and M.M. no. L.11577 ; irregular 
in R.S.M. 1859.33.614 ; replaced by a number of smaller shallow tucks in M.M. no. 
L.10861, and absent in R.S.M. 1859.33.33 (Traquair 1878, pi. 3, fig. i) and other 
specimens. This feature is therefore no more than an individual defect, possibly 
based on an atrophied organ, in the laminar surface layer of the parasphenoid, which 
is frequently wrinkled or irregular when worn (P. 34549 and DMSW.P.149, PL r, 
figs. I, 2) and possibly was denticulated over much of the palatal surface (M.M. no. 
L.11577). 

The hypophysial foramen varies quite obviously in its development, being con- 
spicuous in some specimens (PL i, fig. i) but in others reduced to a very small size 
and accompanied by other diminutive foramina (P. 34556). It was figured without 
comment by both Pander (1858, pi. 3, figs, ir, 13) and Giinther (1871, pi. 34, fig. 4) 
and has been noted by WestoU (1949, text-fig. xD). It is also present in one of the 
specimens of Rhinodipteyus secans, but Gross (1956 : 28, text-fig. 21c) dismissed it as 
" vieUeicht sekundar entstanden ". No such foramen has been recorded from 
Upper Devonian dipnoans. 

The dental plates are continuous with the entopterygoids and appear as mere 
outgrowths of the supporting bones. Fine new specimens of the mandible of the 
earlier Dipnorhynchus from New South Wales (P. 33714, P. 46773) clearly show that 
the rather formless swelhngs that do duty as dental plates are in fact just thickenings 
of the vascular and cosmine layers of the bone of the jaw, and it is evident the more 
definitely patterned " plates " of Dipterus and later dipnoans are of like origin. In 
well preserved specimens they are clearly differentiated by their thick surface layer, 
which has much the same appearance as the cosmine of the external head-bones but 
with rather fewer openings of the fine tubuli, well marked in the low flat longitudinal 
medial areas along which the two plates meet (P . 34543, PL 2, figs, i, 2; R.S.M. 1859 . 
33.33, Traquair 1878, pi. 3, fig. i). In P. 755 and 33166 (PL 2, figs. 3, 4) there is 
irregularity along the symphysial margin due to resorption and in P. 34556 (PL 2, 
fig. 5) as in P. 46691 and R.S.M. 1859.33.612, much of the thick cosmine layer has 
disappeared showing underlying trabecular layer. In extreme cases not only has the 
whole of the surface between the dental plates gone, as in the relatively small original 
of PL I, fig. 2 (DMSW, P. 149), but secondary tubercles [s.d.) may be developed on 
the rough trabecular bone, and an unusual number of denticles appear between the 
regular rows on the biting surfaces (P. 34549, PL i, fig. i. Text-fig. 49). In this 
specimen small irregular areas of the right dental plate (left of figure) have disappeared, 
but this degree of resorption is not seen in any other specimen and must be due to 
decay in an old specimen. Resorption of the plate is not correlated with size for 
the completely resorbed originals of PL i, figs, i and 2 are respectively larger and 
smaller than Traquair 's specimen which is almost perfect. 

The vomerine dentition is not well displayed in any of the available specimens, 



40 



THE HEAD OF DIPTERUS V A L E N C I E N N E S I 



although some show parts of the supporting bone, but one seems to show remains of 
the dentition that can be reasonably interpreted (33166, PI. 2, fig. 4). Watson & 
Day (1916 : 33, text-fig. 6) and others following them showed a pair of elongated 
plates with three transverse rows of small denticles and a narrow hinder end inserted 
medially between the front of the entopterygoid tooth-plates. None of the available 
specimens shows evidence to support this arrangement, which may be based on the 
uneven resorption of the surface of the entopterygoid tooth-plates, as this tends to 
start along the mid-line and in front (P. 34543, P. 755, 33166, PI. 2, figs. 2-4) and the 
narrow extension of these plates is usually broken away. The last of these specimens 
shows a very different arrangement : on each vomer there are two or perhaps three 




Fig. 50. Dipteriis valenciennesi S. & M. Restoration of Skull, palatal view. 



THE HEAD OF DIPTERUS VALENCIENNESI 41 

slightly diverging longitudinal ridges which most probably bore denticles, as Watson 
noted, although there is little evidence of these now. The vomers are well separated 
from the hard " lip " and must have bitten against a cartilage pad in the front of 
the much shorter lower jaw — Giinther's (1871 : 525, pi. 35, figs. 1-3, La.) " lower 
labial cartilage " in Neoceratodus — and their function could have been merely to hold 
before gulping. Giinther (1871 : 518) refers to the " double kind of action " that the 
main upper and lower tooth-plates have to perform, crushing or grinding food on the 
" flat surfaces " and cutting food with the " sharp lateral ridges " which " fit into 
the notches of the opposite tooth like the shells of a Cardium ". So very nicely do 
they fit that there can have been no grinding action involving any degree of lateral 
displacement, so that the movement must be a simple up and down champing, not 
quite vertical, for the lower plate is set at a small angle to the horizontal, sloping 
inwards. And the tooth-plates in Diptenis, with their highly denticulated ridges, 
must have functioned in much the same restricted manner, and such few facets of 
wear as the denticles show bear this out (P . 34549) . Generally speaking the wear is 
due to friction with the food and just blunts the tubercles which fit into pits of their 
own making between the radiating rows of denticles on the opposing plate [e.g., 33166, 
PL 2, fig. 4), and in the lower jaw (P. 46692).^ In old worn dentitions, action was 
much more irregular and new denticles were sometimes developed between the ridges 
(P. 34549, PL I, fig. I). 
A restoration of the palatal aspect of Dipterus is given in Text-fig. 50. 



SUMMARY 

1. The pattern of the dipnoan skull-roof has developed from an ancestral mosaic 
principally by loss and invasion of roofing plates ; fusion occurs for the most 
part as an individual aberration. 

2. The dipnoi were already widely separated from the Crossopterygii (Jarvik 1960 : 
32) at their first appearance, and this separation took place when the skull- 
roof was in a mosaic stage. It is therefore not possible to correlate the plates 
of the dipnoan skuU-roof meaningfully, and attempts to do so must result only 
in geographical approximations for which compound names have no validity. 

3. The use of nomenclature based on that of the skulls of higher vertebrates is 
therefore both misleading and undesirable : a special notation is required and 
a modified edition of Forster-Cooper's alphabetical scheme, properly related to 
the standard sensory canal pattern adequately serves the purpose. 

^ Dr. Gwynne Vevers' observations on the feeding of Neoceratodus at the Zoological Gardens in London 
{in lit. 25 Nov. 64) are very much to the point — " We feed the present specimen on strips of raw horse 
meat or fish approximately 3 X J X J inches. The previous specimens here have all eaten a great deal 
of lettuce, but the present specimen does not appear to enjoy this at all . . . There is no lateral movement 
of the jaws in chewing but a very pronounced up and down nibbling movement along the length of the 
meat strip or fish. The food is then extruded from the mouth and the operation repeated. This may 
happen several times before the food is finally swallowed. These are straight observations. I suppose 
the nibbling movement would be a very suitable way of dealing with fresh water crayfish or a similar 
object." 

In a film recently shown by Mr. I. R. Bishop at the Linnean Society of London (20 Feb. 1965) 
Lepidosiren is seen feeding on live worms with exactly the same action as that described above in 
Neoceratodus. Dipterus may very well have eaten in a similar fashion once food was obtained, but its 
heterocercal tail suggests less sluggish movement than in the living lung-fishes and therefore the possi- 
bility at times of livelier prey. 



42 THE HEAD OF DIPTERUS V A LE N C I E N N E S I 

4. In the early Dipnoi there was an additional row of plates behind the " extra- 
scapular " row of Dipterus (Z-A-Z) which bore the occipital cross commissure 
and was later lost. 

5. Parallel with the relative movement of the roof-bones in Crossopterygii and 
higher vertebrates demonstrated by Westoll (1938), there was a movement 
backwards of the roofing bones of the Dipnoi ; while standard plates were 
developed from the primitive mosaic in front, presumably by loss and inva- 
sion, rows of plates were successively lost at the back of the skuU-roof, the 
occipital cross-commissure being in turn captured by the row in front until 
in Triassic times it was established in YiBY^. After the Devonian period 
there was also an overall reduction in the number of the plates, by invasion 
rather than fusion, a process continued to the present day. 

6. The effect of §2 is further to emphasize the difference between the lung-fishes 
on the one hand and the rhipidistians and coelacanths on the other. It seems, 
indeed, on the evidence of the skull-roof patterns that the Dipnoi split off from 
all the other Osteichthyes at the skull-mosaic stage and developed an 
independent pattern of dermal bones in the skull-roof which has little relevance 
to that of the " Teleostomi " in spite of Westoll's ingenious arguments to the 
contrary and his bold assertion (1949 : 159, text-fig. 10) that " The evidence 
of Devonian forms shows very clearly that the early Dipnoi were very closely 
related to the contemporaneous Rhipidistia " ; my reading of the evidence 
favours very much Jarvik's (i960 : 31-35, text-figs. 28, 30) interpretation of 
their relationships as an isolated offshoot from " Preteleostomi ", and there is 
a lively expectation that the resemblances that they appear to show towards 
the contemporary Rhipidistia will prove of decreasing systematic significance. 
This supposed isolated position is supported by the development of the tooth- 
plates. 

7. In regard to the classification of the Osteichthyes, we are forced back to the ideas 
of the early years of this century, as expressed by Goodrich (1909) when the 
first major cleft in the group was placed between the Dipnoi (without, of course, 
the Arthrodires) and the Teleostomi, equally rated as sub-classes. Lehman 
(1959 : 8) rates the Dipnoi as a Class, but whatever their evaluation, they are 
an isolated group, and the somewhat heated argument as to the significance of 
the term " Crossopterygii " and the appropriate label to cover all the Dipnoi 
Rhipidistia and Coelacanthini (Romer 1955 ; Trewavas, White, Marshall & 
Tucker 1955) is so much beating of the air — the Dipnoi stand apart, and the 
term Crossopterygii covers only the rhipidistians and the coelacanths. 

ACKNOWLEDGMENTS 

Most of the material studied is in the British Museum (Natural History), and to 
the collections there substantial and important additions have been made lately by 
Dr. D. L. Dineley and Mr. John Saxon and his son Alan ; and Mr. Saxon has also 
been able to send valuable information regarding a number of older localities. As 
usual, ready and patient help with invaluable material on loan from the Royal Scottish 
Museum was given by Dr. Charles Waterston, and a number of important fossils 



THE HEAD OF DIPTERUS V A LE N C I EN N E S I 



43 



from the Manchester Museum were generously lent by Dr. R. M. Eagar. These last 
were from the D.M.S. Watson collection, and one from Professor Watson's own col- 
lection was made available for study as well. Dr. F. R. Parrington also kindly sent 
me material on loan from Cambridge. 

In the Department I had my usual full backing by Mr. Harry Toombs, well 
supported in the development of the fossil heads by Mr. Ian Macadie, who also 
influenced the offer of a number of valuable donations from his native Caithness to 
the right quarter. 

The photographs were dexterously taken by Dr. W. T. Dean and Mr. John 
Ferguson, while in putting the final touches to this document Mr. R. H. Spires, Mr. 
F. M. Wonnacott and Mr. R. Baker exercised their respective skills. 

To aU these I offer my sincere thanks. 




: V-.jiK4.x 



Fig. 51. Dipteviis valenciennesi S. & M. Skull-roof of small but aged specimen. Lecto- 
type of " Polyphractus platycephalus " Agassiz (1844 : 5, 29, pi. 27, fig. i). Orkney. 
P-3373«- XI -5. 



44 



THE HEAD OF DIPTERUS V A L E N C I E N N E S I 



REFERENCES 
Agassiz, L. 1833-44. Recherches sur les Poissons fossiles, II. xii + 310 + 336 pp., atlas 

149 pis. Neuchatel. 
■ 1844-45. Monographie des Poissons fossiles du Vieux Gres rouge ou Systeme Devonian 

[Old Red Sandstone) des lies Bvitanniques et de Russia, xxxvi + 171 PP-. atlas 43 pis. Paris. 
Bridge, T. W. 1898. On the Morphology of the Skull in the Paraguayan Lepidosiren and 

in other Dipnoids. Trans. Zool. Soc. Lond., 14 : 325-376, pis. 28, 29. 
FoRSTER-CooPER, C. 1937- The Middle Devonian Fish Fauna of Achanarras. Trans. Roy Soc. 

Edinb., 59 : 223-239, pis. 1-8. 
Goodrich, E. S. 1909. Cyclostomes and Fishes. In Lankester, R. A Treatise on Zoology, 

9, I. 
• 1925. On the Cranial roofing-bones in the Dipnoi. /. Linn. Soc. {Zool.), London, 36 : 

79-86, 7 figs. 

• 1930. Studies on the structure and development of Vertebrates, xxx + 837 pp. London. 

Graham-Smith, W. & Westoll, T. S. 1937. On a new long-headed Dipnoan Fish from the 

Upper Devonian of Scaumenac Bay, P.Q., Canada. Trans. Roy. Soc. Edinb., 59 : 241-266, 

pis. I, 2. 
Greil, a. 1913. In Semon, R. Zoologische Forschungsreisen in Australian und dem Malayis- 

chen Archipel. Denkschr med.- naturw. Ges. Jena, 4, 2. 
Gross, W. 1956. Uber Crossopterygier und Dipnoer aus dem baltischen Oberdevon im 

Zusammenhang einer vergleichenden Untersuchung des Porenkanalsystems palaozoischer 

Agnathen und Fische. K. svenska VetenskAkad. Handl., Stockholm (4) 5, 6 : 1-140, pis. 

1-16. 
GiJNTHER, A. 1871. Description of Ceratodus, a genus of Ganoid Fishes, recently discovered 

in Rivers of Queensland, Australia. Philos. Trans., London, 161 : 511-571, pis. 30-42. 
Hills, E. S. 1933. On a Primitive Dipnoan from the Middle Devonian Rocks of New South 

Wales. Ann. Mag. Nat. Hist., London (10) 11 : 634-644, pis. 11, 12. 
1941- The Cranial Roof of Dipnorhynchus sUssmilchi (Eth. fil.). Rec. Aust. Mus., Sydney, 

21 : 45-55. pl- 9- 

1943- The Ancestry of the Choanichthyes. Aust. J. Set., Sydney, 6 : 21-23, ^S- 

Holmgren, N. & Stensio, E. 1936. Kranium und Visceralskelett der Akranier Cyclostomen 

und Fische. Handb. vergl. Anat., Berlin, 4 : 233-500, 171 figs. 
Huxley, T. H. 1876. Contributions to Morphology. Ichthyopsida, No. i. On Ceratodus 

forsteri, with Observations on the Classification of Fishes. Proc. Zool. Soc. Lond., 1876 : 

24-59, II figs. 
Jarvik, E. 1942. On the Structure of the Snout of Crossopterygians and Lower Gnathostomes 

in General. Zool. Bidr. Uppsala, 21 : 235-675, pis. 1-17. 

1948. On the Morphology and Taxonomy of the Middle Devonian Osteolepid Fishes of 

Scotland. K. svenska VetenskAkad. Handl., Stockholm (3) 25, i : 1-301, pis. 1-37. 

1950. Middle Devonian Vertebrates from Canning Land and Wegeners Halvo (East 

Greenland), II. Crossopterygii. Medd. Grenland, Kobenhavn, 96, 4 : 1-132, pis. 1-24. 

1954- On the visceral skeleton in Eusthenopteron with a discussion of the parasphenoid 

and palatoquadrate in fishes. K. svenska VetenskAkad. Handl., Stockholm (4) 5, i : 1-104, 

47 fig?- 

— i960. Theories de V Evolution des Vertebres. 104 pp., 30 figs. Paris. 

Kesteven, H. L. 1951. The origin of the Tetrapods. Proc. Roy. Soc. Victoria, Melbourne, 

59 : 93-138. 8 figs. 
Lehman, J. -P. 1959. Les Dipneustes du Devonien sup6rieur du Groenland. Medd. Grenland, 

Kobenhavn, 160, 4 : 1-58, pis. 1-2 1. 
Lehmann, W. M. 1956. Dipnorhynchus lehmanni Westoll, ein primitiver Lungenfisch aus dem 

rheinischen Unterdevon. Palaont. Z., Stuttgart, 30 : 21-25, pl. i. 
Lehmann, W. M. & Westoll, T. S. 1952. A primitive dipnoan fish from the Lower Devonian 

of Germany. Proc. Roy. Soc. Lond. (B) 140 : 403-421, pl. 24. 
Marshall, A. J. 1962. In Parker & Haswell " A Text-book of Zoology ". Seventh Edition. 

xxiii -|- 952. London. 



THE HEAD OF DIPTERUS V A LEN C I E N N E S I 45 

Miller, H. 1849. Footprints of the Creator : or the Asterolepis of Stromness. xvi + 313 pp. 

London. 
0RVIG, T. 1961. New Finds of Acanthodians, Arthrodires, Crossopterygians, Ganoids and 

Dipnoans in the Upper Middle Devonian Calcareous Flags (Oberer Plattenkalk) of the 

Bergisch Gladbach — Paffrath Trough, 2. Paldont. Z., Berlin, 35 : 10-27, pls- 1-6. 
Pander, C. H. 1858. Vber die Ctenodipterinen des Devonischen Systems, viii + 64 pp., 9 pis. 

St. Petersburg. 
Parrington, F. R. 1949. A theory of the relations of lateral lines to dermal bones. Proc. 

Zool. Sac. Land., 119 : 65-78, 2 figs. 

1950. The skull of Dipterus. Ann. Mag. Nat. Hist., London (12) 3 : 534-547, 5 figs. 

• 1956. The patterns of dermal bones in primitive vertebrates. Proc. Zool. Soc. Lond., 

127 : 389-411, 9 figs. 
Philip, G. M. & Pedder, A. E. H. 1964. A re-assessment of the age of the Middle Devonian 

of south-eastern Australia. Nature, Lond., 202 : 1323-1324. 
RoMER, A. S. 1936. The Dipnoan Cranial Roof. Amer. J . Sci., New Haven, (5) 32 : 241-256, 

4 figs. 

1955- Herpetichthyes, Amphibioidei, Choanichthyes or Sarcopterygii? Nature, Lond., 

176 : 126. 

1962. Vertebrate Evolution. Copeia, 1962 : 223-227. 

Save-Soderbergh, G. 1932. Preliminary Note on Devonian Stegocephalians from East 
Greenland. Medd. Grenland, Kobenhavn, 94, 7 : 1-107, pis. 1-22. 

1952. On the Skull of Chirodipterus wildungensis Gross, an Upper Devonian Dipnoan 

from Wildungen. K. svenska VetenskAkad. Handl., Stockholm (4) 3, 4 : 1-28, pis. 1-7. 

Sedgwick, A. & Murchison, R. I. 1835. On the Structure and Relations of the Deposits 

contained between the Primary Rocks and the Oolitic Series in the North of Scotland. 

Trans. Geol. Soc. Lond. (2) 3 : 125-160, pis. 13-17. 
Stensio, E. a. 1963. The Brain and the cranial Nerves in fossil, lower craniate Vertebrates. 

Skr. norsk. VidenskAkad. Oslo, I. Mat. Nat. Kl. (N.S.) 13 : 1-120, figs. 1-54. 
Teller, F. 1891. Ueber den Schadel eines fossilen Dipnoers Ceratodus Sturii nov. spec, 

aus den Schichten der oberen Trias der Nordalpen. Abh. geol. Reichsanst. Wien, 15, 3 : 1-38, 

pis. 1-4. 
Traquair, R. H. 1873. On a new genus of Fossil Fish of the order Dipnoi. Geol. Mag., 

London, 10 : 552-554- pl- i- 
1878. On the Genera Dipterus, Sedgw. & Murch., Palaedaphus Van Beneden and De 

Koninck, Holodus, Pander, and Cheirodus, M'Coy. Ann. Mag. Nat. Hist., London (5) 

2 : 1-17, pi. 3. 
Trewavas, E., White, E. L, Marshall, N. B. & Tucker, D. W. 1955. [Discussion of A. S. 

Romer 1955]. Nature, Lond., 176 : 126. 
Wade, R. T. 1935. The Triassic Fishes of Brookvale, New South Wales, xiv -\- 89 pp., 

10 pis. British Museum (Nat. Hist.), London. 
Watson, D. M. S. & Day, H. 1916. Notes on some Palaeozoic Fishes. Mem. Manchr. Lit. 

Phil. Soc, 60, 2 : 1-52, pis. 1-3. 
Watson, D. M. S. & Gill, E. L. 1923. The structure of certain Palaeozoic Dipnoi. J . Linn. 

Soc. {Zool.) London, 35 : 163-216, 34 figs. 
Westoll, T. S. 1938. Ancestry of the Tetrapods. Nature Lond., 141 : 127-128, 2 figs. 

1943- The Origin of the Tetrapods. Biol. Rev., Cambridge, 18 : 78-98, 9 figs. 

1944- The Haplolepidae, a new Family of late Carboniferous Bony Fishes. Bull. Amer. 

Mus. Nat. Hist., New York, 83, i : 1-122, pis. i-io. 

1949- On the Evolution of the Dipnoi. In Genetics, Paleontology, and Evolution (Ed. 

Jepsen, G. L., Simpson, G. G. & Mayr, E.) : 1 21-184, ^i ^gs. Princeton. 

White, E. I. 1962. A Dipnoan from the Assise de Mazy of Hingeon. Bull. Inst. roy. Sci. nat. 

Belgiqiie, Bruxelles, 38, 50 : 1-8, pis. i, 2. 
Woodward, A. S. 1891. Catalogue of the Fossil Fishes in the British Museum [Natural History), 

2. xliv -\- 567 pp., 16 pis. British Museum (Nat. Hist.), London. 



PLATE I 
Dipterus valenciennesi S. & M. 

Fig. I. Undersurface of skull of old specimen showing decay (or irregular resorption) of right 
tooth-plate (left of figure) and complete resorption of surface of entopterygoid between tooth- 
plates with formation of secondary denticles. CoUd., D. L. Dineley 1957. Clardon Haven, 
Caithness. P. 34549. X2. See also Text-fig. 49. 

Fig. 2. Undersurface of skull showing vertebral elements fused to basioccipital region. Loo. 
unknown, Caithness. DMSW.P.149. X2. 



Bull.B.M. [N.H.) Geol. ii, i 



PLATE 1 




PLATE 2 

Dipterus valenciennesi S. & M. 

Fig. I. Imperfect entopterygoid tooth-plates showing cosmine surface and hypophysial 

foramen. Colld., D. L. Dineley, 1957. Clardon Haven, Caithness. P. 34543. X3. 

Fig. 2. Same specimen powdered to show initial resorption along median suture, x i -5. 

Fig 3. Another specimen showing increased resorption along median suture. Orkney. 

P. 755- X 1-5 

Fig. 4. Specimen showing advanced and irregular resorption along median suture and worn 
vomerine dentition in front. Near Thurso, Caithness. 33166. x 1-5. 

Fig. 5. Palatal view of skull. The median area of the entopterygoid tooth-plates has lost 
the cosmine surface completely, and only the base of the vomerine dentition remains. Colld., 
D. L.Dineley, 1957. Clardon Haven, Caithness. P. 34556. xi-5. 



Btill. B.M. {N.H.) Geol. ii, i 



PLATE 2 




PLATE 3 
Dipterus valenciennesi S. & M. 

Fig. I. Skull-roof showing pores of sensory canals enlarged by resorption and the post- 
cranial processes (" tabular horns "). Colld. J. Saxon, 1964. Weydale Quarry, Caithness. 
P. 46761. X2. 

Fig. 2. Hinder end of same specimen. The left post-cranial process and plate Ij have lost 
outer and part of middle layers exposing section of occipital cross-commissure with short branch 
forwards. X4-5. 

Fig. 3. End of snout showing pores of supraoccipital canal enlarged by resorption. (See also 
Text-fig. 19) Colld. R. V. Collier, 1964. Clardon Haven, Caithness. P. 46690. X4*3. 



Bull. B.M. (N.H.) Geol. ii, i 



PLATE 3 




-•V,,j^ 




I'^^T^'-S'iV 




















% 

196* 



PRINTED IN GREAT BRITAIN 
BY ADLARD & SON LIMITED 
BARTHOLOMEW PRESS, DORKING 






SOME LOWER 



CRETACEOUS TEREBRATELLOIDEA 



E. F. OWEN 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. ii No. 2 

LONDON: 1965 



SOME LOWER -^ ^ 

CRETACEOUS TEREBRATELLOIDEA \% 




BY 

ELLIS FREDERIC OWEN 



-?<_ 



Pp. 47-72 ; 3 Plates ; 13 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

GEOLOGY Vol. 11 No. 2 

LONDON: 1965 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 2 of the Geological 
(Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
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TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 7 September, 1965 Price £1 



SOME LOWER 
CRETACEOUS TEREBRATELLOIDEA 

By ELLIS FREDERIC OWEN 

SYNOPSIS 

Several Lower Cretaceous species previously described by Walker, Davidson, Meyer and 
Keeping are revised and referred to Tamarella gen. nov. or Vectella gen. nov. Serial sections of 
representative species of these forms are shown for the first time. In addition, two new species 
of Rugitela from the Neocomian of Lincolnshire and Yorkshire are described and related to forms 
previously recorded from North Germany and France. 

INTRODUCTION 

Representative genera of the Mesozoic brachiopod family Zeilleriidae have been 
recorded from the Trias to the Lower Cretaceous but few of the early records have 
been further investigated, and the Jurassic genus Zeilleria has often been quoted from 
the Neocomian and, more recently, with those of Aulacothyris and Ornithella by 
Middlemiss (1959) and Casey (1961), from Aptian and Albian beds of the Lower 
Greensand. Some of these Cretaceous brachiopods, while possibly belonging to the 
same families as Jurassic species, have characters which merit generic distinction. 

Until the researches of Muir-Wood (1934, 1936) the internal structures of the British 
Jurassic Terebratelloidea had been known only from dissections or by chance 
exposure of brachial loops and cardinalia in damaged specimens. Minor differences 
in loop development had previously escaped critical investigation and no real 
advance had been made in their further classification. It is not the purpose of this 
paper to add to or further discuss the classification of the terebratelloid brachiopods. 
However, a systematic study has been made of some of the Lower Cretaceous species 
within this suborder together with comparisons of genera already described by Muir- 
Wood (1934, 1936) and Cooper (1955). 

Some of the early Cretaceous terebratelloid species formerly described by Walker 
(1867, 1868, 1870) from the Lower Greensand of Upware (Cambridgeshire), Brickhill 
(Buckinghamshire) and Potton (Bedfordshire) have been revised and referred to 
Tamarella gen. nov. Others, which had been broadly interpreted as Waldheimia, 
Zeilleria, Aulacothyris and Ornithella are placed in Vectella gen. nov. 

Keeping (1883 : 24) suggested a possible line of evolution of some of Walker's 
species and related them to others described by Morris (1854) and J. de C. Sowerby 
(1836) from the Lower Greensand of the Isle of Wight. His suggestions were, 
however, based mainly on a comparison of external morphology, and an examination 
of the species listed by him has revealed a degree of homoemorphy. Information 
obtained from transverse serial sections of these forms has now made possible a 
revision of his ideas regarding their relationships. 

In a description of a brachiopod fauna from the Mural Limestone (Middle Albian) 
of Arizona, Cooper (1955 : 10) referred Terehratnla tamarindiis J. de C. Sowerby to his 
new genus Psilothyris. He figured a series of etched specimens showing (pi. 3) what 

5§ 



50 



SOME LOWER CRETACEOUS TE RE BR ATE LLO 1 DE A 



he described as precampagiform, terebrataliform and dalliniform stages in the 
development of the brachial loop of certain specimens. On the same plate, for 
comparison, he figured the cardinalia without brachial loop of a dissected specimen 
which he identified as T. tamarindus from the Lower Greensand of Faringdon, 
Berkshire. If, as he suggested, the etched loops of his specimens exhibit dallinoid 
development stages, then Psilothyris would appear to belong wdth the Dallinidae. 





D 




Fig. I. Diagram illustrating the cardinalia of (A) Taniarella, showing the broad hinge- 
plates with the anterior plication, and (B) a reconstruction of the brachial loop showing 
the thickening of the descending branches. (C) Vectella, showing the deep hinge-trough 
and long dorsal septum, and (D) reconstruction of the brachial loop. 

Although many young specimens of T. tamarindus have been examined from the 
Lower Greensand of Faringdon, Berkshire, no dallinoid stages in loop development 
have been found, but a remarkable specimen of T. tamarindus from Faringdon 
(PL I, fig. 8c) shows what appears to be a thickening of the descending branches some 
6 mm. below the dorsal umbo in a brachial valve, 15 mm. long. It is possible that 
this thickening may be the remains of an early connecting band or attachment. A 
similar thickening is seen on the descending branches of the brachial loops of speci- 
mens of Psilothyris figured by Cooper (1955, pi. 3, figs. 18, 19). 

The lack of serial sections of the genus Psilothyris make absolute comparison with 
British Upper Aptian species of Tamarella impossible and the absence of any evidence 
of early loop attachment in even very young specimens of Tamarella means that any 
attempt to relate the two genera is purely conjecture. However, the two genera have 
much in common and it may be necessary at some future date to emend the present 
study. 

The varied brachiopod fauna contained in the condensed Neocomian beds represen- 
ted in the Claxby Ironstone of Lincolnshire and the Speeton Clay of Yorkshire is 



SOME LOWER CRETACEOUS TERE BR ATELLO IDE A 51 

comparable to that described by Rocmcr (1836-1840) from the Hilsconglomerat of the 
Hanover district, north-west Germany. Species of zeilleriid brachiopods described as 
Terebratiila hippoptis and T. longa by Roemer are revised here and assigned to the 
genus Rugitela Muir-Wood from the Fuller's Earth Rock of Somerset and said at the 
time to have a probable range of ? Inferior Oolite to ? Upper Jurassic. 

Judd (1868) was the first to recognize Roemer's species T. hippopus in Lincolnshire 
where it occurs fairlj^ abundantly in the Claxbj' Ironstone. Davidson (1874) erected 
a varietal name for the British form on the grounds of its larger dimensions, but these 
are directly proportionate to specimens from the type locality at Berklingen and 
from Saltzgitter, near Hanover. Furthermore, the pattern of variation in outline, 
width of shell and depth of dorsal sulcus is identical in both forms and there seems no 
point in upholding Davidson's varietal name. 

SYSTEMATIC DESCRIPTIONS 

Family ZEILLERIIDAE Allan, 1940 

Genus VECTELLA nov. 

Diagnosis. Shell elongate-oval, sulco-carinate to biconvex, rectimarginate to 
uniplicate or sulcate. Umbo massive, beak suberect. Foramen circular. Beak- 
ridges sharp, mesothyrid ; interarea short. Test smooth, finely punctate. Delti- 
dial plates conjunct. Short, subparallel dental lamellae embedded in callus ; in 
mature individuals these lamellae support massive, peg-like, inwardly directed hinge- 
teeth. Cardinal process poorly developed. Acute septalium forms broad V-shaped 
hinge-trough supported by short, thick, persistent dorsal median septum. Crural 
bases triangular, giving rise to zeilleriiform brachial loop developed ventrally. 

Type species. Waldheimia celtica Morris 1854 : 158. 

Locality and horizon. In addition to the type-species, which comes from the 
Upper Aptian, Parahoplites nutfieldensis Zone, Shanklin, Isle of Wight, Vectella is 
represented at an equivalent horizon at Upware (Cambridgeshire) by V. woodwardi 
(Walker) and V. angusta (Walker) and by V. morrisi (Meyer) from the Bargate Beds 
of Surrey, from Brickhill (Buckinghamshire) and from Shanklin, Isle of Wight. 

Remarks. Vectella is probably a further development of the Jurassic genus 
Ornithella but differs in its more acute septalium and deeper hinge-trough, more 
acutely triangular hinge-plates and crural bases, and more extensive inner socket- 
ridges. It differs from Tamarella nov. and Rugitela in its thicker shell, fused dental 
lamellae and narrower, anteriorly plane, septalium. 

Vectella celtica (Morris) 
(PI. I, figs, ^a-c ; Text-fig. 2) 

1847 Terebratula longa Roemer ; Davidson & Morris : 255, pi. 19, figs, i & za-d. 

1854 Waldheimia celtica Morris : 158. 

1855 Waldheimia {Terebratula) celtica Morris ; Davidson : 73, pi. 9, figs. 32-35. 
1874 Waldheimia celtica Morris ; Davidson : 47, pi. 6, fig. 15. 

Description. Biconvex, elongate-oval zeilleriid brachiopod, approximately 



52 some lower cretaceous tere br atello i d e a 

0-6 0-2 ^^^01 0-4 ^PP W|||^ 



r\r\ 



0-3 



( 1 





0-4 



Fig. 2. A series of sixteen transverse sections through the umbonal part of Vectella celtica fMorris) 
from the Upper Aptian, ShankHn, Isle of Wight. BM. B. 25801. X2. 

30 mm. long, 19 mm. wide and 18 mm. thick. It has a short, massive umbo, sub- 
erect beak and well exposed symphytium. The circular foramen is fairly large with 
mesothyrid beak-ridges bordering a short interarea. In young forms a marked 
sulcus is developed posteriorly in the brachial valve and may still be seen in some 
specimens of mature forms. 

Lectotype. Morris was the first to use the specific name Waldheimia celtica for a 
brachiopod from the Lower Greensand of Great Britain and listed it in the second 
edition of his catalogue (Morris 1854 • 158)- He illustrated the species by indicating 
specimens in Davidson's Cretaceous Monograph which was not, in fact, published until 
a year later (Davidson 1855 : 73, pi. 9, figs. 32-35). One of these specimens had 
certainly been taken from a series of four given to Davidson by Morris and now in the 
Davidson Collection at the British Museum (Natural History), registered number 
B • 6757. The specimens are here regarded as syntypes of V. celtica (Morris) and one, 
figured by Davidson (1855, pi. 9, fig. 33) is chosen as lectotype and has been re- 
registered BB. 42915. 



SOME LOWER CRETACEOUS TE RE B R A T E LLO I D E A 53 

Material. The lectotype and numerous specimens in the Davidson Collection 
and general collection of the British Museum (Natural History) from the Upper 
Aptian, Parahoplites nutfieldensis Zone of Shanklin, Isle of Wight, registered as 
B . 6757, B . 25800, B . 25802-25810. A specimen, B . 14728, in the Sedgwick Museum, 
Cambridge shows the brachial loop and cardinalia in an almost perfect state of 
preservation. 

Remarks. This species probably illustrates an evolutionary link with some 
species of the Jurassic genus Ornithella. Externallj', the long tapering oval outline, 
massive umbo and carinate pedicle valve suggest such species as 0. ornithocephala 
(J. Sowerby) and 0. bathonica (Rollier). Internally the thickened valve walls and 
callus embedded dental lamellae in the pedicle valve and the thick septum, sup- 
porting short, stout hinge-plates, have much in common with those of 0. bathonica 
as shown in transverse serial sections by Muir-Wood (1934 : 544). It can be dis- 
tinguished from either of the two Jurassic species in its shorter umbo, larger foramen 
and less acutely tapering valves. It is distinguished from other Lower Cretaceous 
forms, such as V. woodwardi (Walker), by its more regular oval outline, more in- 
curved beak, smaller foramen and absence of anterior sulcation of the brachial valve. 
It differs from V. morrisi (Meyer) in its greater dimensions, stronger biconvexity and 
lack of uniplication of the anterior margin. It bears a superficial resemblance to 
V. angusta (Walker) from Upware and Brickhill but differs in having a more incurved 
beak and more robust general outlines. It is nevertheless very near this species. 

Vectella angusta (Walker) 
(PI. I, figs. 5a-c) 

1868 W aldhehnia niiitabilis var. angusta Walker : 400, pi. 19, fig. 5, 5a. 

1870 Waldheimia mittabilis var. angusta Walker ; Walker : 562. 

1874 Waldheimia wanklyni var. angusta Walker ; Davidson : 51, pi. 7, figs. 26-28. 

Description. Elongate-oval to fusiform Vectella about 34 mm. long, 18 mm. 
wide and 12 mm. thick. Maximum width midway between umbo and anterior 
margin. Biconvex with acute carination of pedicle valve. Large foramen domina- 
ting a slightly produced suberect umbo with wide interarea and well exposed sym- 
phitium. Extensive beak-ridges are well defined. Anterior commissure plane. 

HoLOTYPE. British Museum (Natural History) no. 67601, figured by Walker 
(1868, pi. 19, figs. 5, 5a) as Waldheimia mittabilis var. angusta. From the Lower 
Greensand of Upware, Cambridgeshire. 

Material. In addition to the holotype, eight specimens from Upware, registered 
no. B.2712 and seventeen specimens from Brickhill, Buckinghamshire, registered no. 
B. 25503, all in the British Museum (Natural History). 

Remarks. As in the case of Tamarella elliptica (Walker) , this species was originally 
described as a variety of Waldheimia mutabilis, a name already used by Oppel (1861 : 
538) for a Liassic species. On Walker's instructions the name W. itianklyni was 
substituted by Davidson (1874 : 51) for W . mutabilis Walker. 

Walker neither selected nor indicated a type specimen for W. mutabilis [= W. 
wanklyni] and the species is, therefore, in doubt. The variety angusta, however, is 

5§§ 



54 SOME LOWER CRETACEOUS TE R E B R A T E LLO 1 IJ E A 

easily recognizable as a form from the Lower Greensand of Upware and Brickhill. 
It is in no way related to the species Tamarella elliptica and is here promoted to specific 
rank. 

Somewhat resembling V . celtica (Morris) in general outline, V. aiignsta differs in less 
clearly defined features, more acutely convex brachial valve and produced umbo. 
The form from Brickhill has a shorter umbo and is more regularly elongate-oval in 
outhne than the typical form. It also has a slight depression or sulcus visible on the 
brachial umbonal region. This depression is more marked in younger specimens. 

Vectella woodwardi (Walker) 
(PI. I, fig. la-c ; Text -fig. 3) 

1867 Waldheimia woodwardi Walker ; 455, pi. 19, fig. 3. 

1868 Waldheimia woodwardi Walker ; Walker : 404. 

1874 Waldheimia woodwardi Walker ; Davidson : 52, pi. 6, figs. i-5rt. 
1883 Waldheimia woodwardi Walker ; Keeping : 21. 

Description. Elongate-oval Vectella, about 37 mm. long, 20 mm. wide and 19 
mm. thick. The pedicle valve is acutely carinate with steep flanks. It has a short, 
suberect umbo truncated by a large, circular foramen. Sharp mesothyrid beak- 
ridges border a short, flat interarea. The finely punctate shell surface is covered by 
numerous fine growth-lines. A shallow sulcus originating from the umbo in the brachial 
valve broadens anteriorly. The internal characters are as described for the genus. 

Lectotype. The original specimen described and figured by Walker (1867) is one 
of the two syntypes originall}' registered with the Walker Collection in the British 
Museum (Natural History) as 62202. It has been re-registered as BB. 42910 and is 
here selected as lectotype. 

Material. Eighteen specimens in the general collection of the British Museum 
(Natural History) nos. B. 25717, B. 25801, 62202, BB. 21130-38, BB. 42922, all from 
the type locality at Upware, and several internal moulds from Potton. 

Remarks. Vectella woodwardi is known from the type locality at Upware (Cam- 
bridgeshire) and from Potton (Bedfordshire), where it occurs in the Parahoplites 
mttfieldensis Zone of the Upper Aptian. In many respects it resembles V. celtica 
(Morris) which occurs at the same horizon at Shanklin, Isle of Wight but differs in 
shell convexity and beak characters. These differences in morpholog}^ may be due 
merely to change of environment. 

The broad sulcation of the brachial valve in V . woodwardi and the shorter, more 
massive umbo and larger foramen distinguish the species from V . morrisi (Meyer) and 
V. angitsta (Walker). 

Vectella morrisi (Meyer) 

(PI. I, fig. 7 ; Text-fig. 4) 

(PI. 3, figs. 7-9) 

1863 Terebralula moiUoniana Ltmkester : 414, pi. ig, figs. 1-3. (uon d'Orbigny 1848). 

1864 Waldheimia moitioniana Lankester ; Meyer : 231, figs. 12-14. 
1868 Waldheimia morrisi Meyer : 269. 

1874 Waldheimia morrisi Mej'er ; Davidson : 47. 



SOME LOWER CRETACEOUS T E RE BR AT li LLO 1 D E A 

0-4 0-2 0-2 0-3 W 0-5 ^BP^^, 



ooact 



55 



07 0-4 ^^^ 0-3 ^^^ 0-3 

mm m m O^ 




Fig. 3. Sixteen tninsversc serial sections througli the umbo of Vcctella woodwardi (NValker) from 
the Upper Aptian, Upware, Cambridgeshire. BM. B. 42922. X2. 

Description. Although somewhat terebratulid in general aspect, Vectdla 
morrisi is characteristically oval in outline with a shorter, more incurved umbo than 
other described species of Vcctella. Both valves are comparatively flat but the 
brachial valve has a marked median ridge extending two-thirds of the shell length 
and giving rise to fairly steep flanks. The pedicle valve remains gently convex. 
The anterior commissure is marked by a faint uniplication distinguishing it from 



56 



SOME LOWER CRETACEOUS TE R E BR ATE LLO I D E A 



V. celtica (Morris) which it somewhat resembles. The typical form occurring at 
Shanklin, Isle of Wight reaches a maximum length of 23 mm., a width of 17 mm. and 
attains an average thickness of 11 mm. 

Lectotype. Meyer (1864 : 249) first described the species as Terebratula moutoni- 
ana Lancaster and later (1868 : 269) re-described it under the binomen Waldheiniia 
morrisi referring to the figured specimens (1864 : figs. 12-14) of his earlier work. 
Two of the specimens figured by Meyer (figs. 12, 13) were stated to have been collected 
from the Pebble-bed of Shanklin, while the other (fig. 14) was said to have come from 
the Pebble-bed of Godalming, Surrey. 

These specimens, here regarded as syntypes of V. morrisi, are in the Sedgwick 
Museum, Cambridge and are registered as B . 14770-71 and B . 16785. The lectotype, 
here selected, is B.14771. It was originally figured by Meyer (1864, pi. 12, fig. 13) 
and was collected from the Pebble-bed, Parahoplites nutfieldensis Zone, Shanklin, Isle 



of Wight. 



o 




Fig. 



Fourteen transverse serial sections througli tlie umbo of Vcctella morrisi (Mej'er) from 
the Upper Aptian, Shanklin, Isle of Wight. BM. B. 2 1937. X2. 



Material. Apart from the type specimens mentioned above, there are forty-two 
specimens in the Sedgwick Museum, Cambridge, twenty-nine from Shanklin and 
thirteen from the Bargate Pebble-bed of Godalming, Surrey, registered B. 16838-50, 
B. 14761-69, B. 14776-89. Also there are fifty-four specimens in the general collec- 
tion and Davidson Collection of the British Museum (Natural History) registered 
B. 25506-9, B.6740, B. 25815-16, BB. 42914, BB. 42916-18. 

Remarks. A smaller form of V . morrisi, though with similar proportions, occurs 
in the Bargate Pebble-beds of Surrey and beds of equivalent age at Brickhill, Bucking- 
hamshire. The latter has often been confused with TamareUa juddi (Walker) but 
differs from this species in its more regular oval outline, less acutely convex valves, 
steeper flanks, shorter umbo, less extensive interarea, rounded beak-ridges and 
simpler zeilleriform brachial loop. It is approximately 17 mm. long, 11 mm. wide 
and 8 mm. thick. 



SOME L0\VI':R cretaceous TEREBRATEELOIDEA 57 

Genus TAMARELLA no v. 

Diagnosis. Shell biconvex. Circular to elongate-oval to pentagonal in outline. 
Folding rectimarginate to incipiently uniplicate to ligate. Umbo massive, suberect ; 
beak-ridges sharp. Deltidial plates conjunct. Foramen large, mesothyrid. Shell 
surface often with marked concentric growth-lines. Median septum short, extending 
less than one third the length of brachial valve. Septalium broad, shallow, anteriorly 
arched. Hinge-plates fused, overlapping. Long brachial loop unattached to median 
septum in adult stage, given off ventrally. 

Type species. Terebratula tamarindus J. de C. Sowerby 1836. 

Range. Upper Aptian. 

07 0-2 ^01 •^O^ M^O-3 >i^0-2 >Mfc^( 



0-4 







Fig. 5. Twelve transverse serial sections through the umbonal part of Tamarella tamarindus 
(J. de C. Sowerby) from the Upper Aptian, Shanklin, Isle of Wight. BM. B.506. X2. 



Tamarella tamarindus (J. de C. Sowerby) 

(PI. I, figs. 2a-c, 8a-c, loa-c, PI. 3, ligs. ^a-c, 6a-c ; Text-figs. 5, 6) 

1836 Terebratula tamarindtis J. de C. Sowerby : 338, pi. 14, fig. 8. 

1843 Terabratula tamarindus Sowerby ; Morris : 137. 

1855 W aldheimia tamarindus (J. de. C. Sowerby), Davidson : 74, pi. 9, figs. 26-31. 

1868 Terebratula tamarindus var. magna Walker : 465, pi. 19, fig. 10. 

1874 Waldheimia tamarindus var. magna (Walker) Davidson : 49, pi. 6, figs. 16-19^. 

1955 Psilothyris tamarinda (Sowerby) Cooper : 14, pi. 3, fig. 25. 

Description. Tamarella subcircular to elongate-oval in general outline. Acutely 
biconvex to almost orbicular. Approximately 11 mm. long, 10 mm. wide and 8 mm. 
thick. The massive umbo is truncated by a large foramen. Well marked meso- 
thyrid beak-ridges border extensive interarea. Shell surface evenly punctate and 
ornamented by fine concentric growth-lines. The anterior commissure is rectimar- 
ginate to incipiently uniplicate. There is sometimes a shallow anterior depression or 
sulcus bordered by faint carinae noticeable on the pedicle valve. Shell margins show 
a tendency to gerontic thickening. 

Neotype. J. de C. Sowerby in Fitton (1836 : 338) described a zeilleriid brachiopod 
from the Lower Greensand near Hythe, Kent, under the name Terebratula tamarindus. 
No type material was indicated but a specimen, stated to belong to the Sowerby 



5S 



SOME LOWER CRETACEOUS TE R E B R A TE LLO I DE A 



Collection, was figured (pi. 14, fig. 8) and is now lost. Sowerby's description refers to 
a specimen with an orbicular outline and with beak-ridges extending some distance 
down the sides of the valves. The figured specimen appears flattened in lateral 
profile and with short beak-ridges which are not clearly defined. Although its geo- 



0000 

0-6 ^ 0-2 ^P^ 0-4 ^IP^ 




Fig. 6. Seventeen transverse serial sections through the umbo of Tamarella tamarindus (J. de C. 
Sowerby) from the Upper Aptian, Upware, Cambridgeshire. BM. B. 25693. X2. 



SOME LOWER CRETACEOUS TE R E B R ATE LLO I DE A 59 

logical age was stated to be Lower Greensand, no precise beds were mentioned 
within this horizon nor was any associated fauna listed. Its geological age must, 
therefore, remain broadly defined. Furthermore, the type locality, said to be "near 
Hythe, Kent ", although suggesting a Hythe Beds origin, might alternatively have 
indicated the Sandgate Beds (Upper Aptian) or even an Albian horizon. 

The present widelj^ accepted connotation of a brachiopod under the specific name 
tamarindus is Davidson's description (1855 : 74) under the binomen Waldheiniia 
tamarindtis (J. de C. Sowerby). Davidson figured six specimens from various 
localities of Lower Greensand age, including one specimen (pi. g, fig. 26) stated to 
have been collected from between Hythe and Sandgate, Kent. Unfortunately none 
of the six specimens figured by Davidson is available in the Davidson Collection and 
they cannot be traced elsewhere. 

Although Sowerby's definition of Terebratula tamarindus was vague, the name is so 
entrenched in literature that it would seem a pity not to preserve it. The preserva- 
tion of the name, however, depends on the erection of a neotype to replace Sowerby's 
original figured specimen. 

The Lower Greensand includes, among other horizons, the Sandgate Beds (Upper 
Aptian), which crop out between Hythe and Sandgate, Kent, and from which 
Sowerby's specimen is likely to have been collected. It is no longer possible to 
collect brachiopod specimens from the Sandgate Beds exposed in this area but beds of 
an equivalent age falling within the Parahoplites nutfieldensis Zone occur at Shanklin, 
Isle of Wight and occasionally contain specimens which agree with the original 
description of Terebratula tamarindus J. de C. Sowerby. They also compare 
favourably with specimens figured by Davidson (1855) from " near Sandgate " and 
from the Isle of Wight. 

A specimen probably used by Davidson (1855 : 74) in his description of the species 
is figured here (PI. 3, fig. 5) and proposed as neotype. This specimen, originally 
registered as B . 6724, forms part of the Davidson Collection in the British Museum 
(Natural History) and was collected from the LTpper Aptian beds of Shanklin, Isle of 
Wight. It has been re-registered as BB. 42907. 

Material. A further sixty-one specimens of Tamarella tamarindus (J. de C. 
Sowerby), all from the Isle of Wight, are contain in the general collection and the 
Davidson Collection at the British Museum (Natural History) and are registered as 
B,5o6, B.6724, B. 25820, B. 25187, BB. 42905, BB. 42906, BB. 42919, BB. 42920. 

Remarks. A larger form of this species approximately 24 mm. long, 21 mm. wide 
and 15 mm. thick occurs in the Upper Aptian at Upware and Potton. It was 
originally described and figured as Terebratula tamarindus var. magna by Walker 
(1868 : 465, pi. 19, fig. 10) but agrees in every detail with the typical form and has 
proportionately similar dimensions. T. tamarindus is distinguished from other 
described species of Tamarella by its acute biconvexity and almost circular general 
outline. 

Tamarella bonneyi (Keeping) 
(PI. I, figs. 3a-c ; Text-fig. 7) 
1883 Waldheimia bonneyi Keeping : 129, pi. 7, fig. 4. 



6o 



SOMIi L(nVEK CRKTACliOUS TE RE B R AT E I. LO 1 1) E A 



Description. Oval to pentagonal, somewhat cinctiform in outline, this species 
attains an approximate length of 32 mm. with a maximum width of 24 mm. and 
thickness of 17 mm. Each broad, flattened valve has a shallow anterior sulcation 
and is ornamented by numerous distinct or clearly defined concentric growth-lines. 
The umbo is not so produced as in other species of Tamarella but shows the samebroad, 
flat interarea bordered by sharp beak-ridges. Although constantly biconvex, the 
degree of convexity of each valve varies considerably in T. honneyi. Some forms 
have an almost flat pedicle valve with an acutely convex brachial valve. Others are 
almost equally biconvex and, apart from their greater length, resemble T. tamarindiis 
from Upware. 



e <f> O O <0> 



00 




Fig. 7. A series of fourteen transverse serial sections through the umbo of Tamarella bonneyi 
(Keeping) from the Upper Aptian of Brickhill, Buckinghamshire. BM. B.254Q3. X2. 

HoLOTYPE. B. 26803 from the Lower Greensand, Upper Aptian of Brickhill, 
Buckinghamshire in the collections of the Sedgwick Museum, Cambridge. 

Material. In addition to the holotype, thirty-eight specimens from Brickhill 
are in the Sedgwick Museum registered as B. 80836-39, B. 26509-19, B. 25490-5 13 
and thirty-six specimens in the general collections of the British Museum (Natural 
History) registered as B . 25492-93, B . 25499. 

Remarks. There is a marked similarity between this species and Tamarella vesta 
sp. nov. and attempts have been made to illustrate gradation from one species to 



S().MI<: LOWER CKIiTACEOUS T E R E B R A T !■; L LO I I) !•: A 



f)i 



another. A series registered B. 25490-531 in the Sedgwick Museum illustrates this 
variation. T. bonneyi, however, appears to be confined to the deposits at Brickhill 
and T. vesta is only rarely found at this locality while it is abundant at Upware. 

Tamarella bonneyi is distinguished from T. vesta and other species of Tamarella 
mainly by its larger dimensions, broad cinctiform outline and more marked concentric 
growth-lines. 

Tamarella juddi (Walker) 
(PI. I, figs, iia-c ; Text -fig. 8) 

1868 Waldheimia rhomboidea Walker : 400, pi. 18, fig.s. 3, 4. 

1870 Waldheimia juddi Walker : 562. 

1874 Waldheimia juddi Walker; Davidson : 50, pi. 7, figs. 15-18. 

Description. Tamarella about 19 mm. long, 14 mm. wide and 12 mm. thick. 
Elongate-oval in outline with a maximum width just less than half the distance from 
the umbo to the anterior margin. Acutely biconvex in lateral profile with marked 
marginal growth-lines. Short massive umbo and suberect beak with large foramen 
and sharp mesothyrid beak- ridges. An extensive interarea exposes conjunct deltidial 
plates. Anterior margin laterally compressed and incipiently uniplicate. 

Lectotype. Although the species was originally described by Walker (1868 : 
400) as Waldheimia rhomboidea, it has never been referred to under this name. 
Walker mistakenly thought the name to be preoccupied by Terebratula rhomboidea 
Barondi 1855, from the Tertiary of Italy and subsequently changed the name (1870 ; 
562) to Waldheimia juddi. As a junior synonym of W. rhomboidea, W . juddi has 

1-2 0-4 0-6 0-2 ^0-5 Mt^. 







Fig. 8. Thirteen transverse serial sections through the umbo of Tamarella juddi (Walker) from 
the Upper Aptian of Upware, Cambridgeshire. BM. B. 25681. X2. 



6z SOME LOWER CRETACEOUS T E R E BR AT E LLO I D E A 

since become entrenched in literature as the name of a well known fossil brachiopod 
and is maintained here. Walker's (1868) original description was accompanied by 
figures of two specimens from the type locality at Upware which are in the general 
collections at the British Museum (Natural History) registered as 67602. The larger 
has been re-registered as BB .42929 and is here selected as lectotype (PL i, figs, iia-c). 

Material and locality. In addition to the type material, there are fifty-eight 
specimens in the British Museum (Natural History) registered as B. 25681 and 
B. 25682. Among seven specimens in the Sedgwick Museum from the Upper Aptian 
of Faringdon, registered as B. 18400-06, one specimen, B. 18400, shows a well 
exposed brachial loop and cardinalia which appear to be very similar to Cooper's 
figure of Psilothyris (Cooper 1955, pi. 3, fig. 19). The thickened portion of the 
descending branches can be clearly seen. 

Remarks. Tamarella jiiddi is distinguished from other species of Tamarclla by 
its acute biconvexity, extensive interarea, laterally compressed and tapering anterior 
and smaller foramen. 

Tamarella vesta sp. n. 

(PI. I, figs, ga-c ; Text-fig. 9) 

1868 Waldheimia pseiidojiivensis (Leymerie) ; Walker : 405, pi. 18, figs. 8-1 1. 
1874 Waldheimia pseudojurensis (Leymerie) : Davidson : 48, pi. 7, figs. 12-14. 

Diagnosis. Tamarella about 22 mm. long, 15 mm. wide and 10 mm. thick. 
Shell biconvex, elongate-oval to pentagonal in outhne. Anterior commissure plane. 
Folding ligate. Test smooth with prominent growth-lines. Umbo slightly produced, 
suberect. Deltidial plates conjunct, well exposed. Beak-ridges sharp. 

Description. Although characteristically elongate with steep flanks and slightly 
produced umbo, the range of variation within this species includes forms which are 
broader and, in some cases, more inflated than the typical. Marked gerontic thicken- 
ing of the shell margins tends to accentuate the anterior sulcation of each valve. 

The generic characters seen in transverse serial sections (Text-fig. 9) show very 
clearly the overlapping hinge-plates which appear to be more marked in T. vesta 
than in the type-species T. tamarindtis (J. de C. Sowerby). 

HoLOTYPE. This was originally one of four specimens in the Walker Collection, 
British Museum (Natural History) which were used in Walker's description of the 
species under the name Waldheimia pseudojurensis Leymerie. The holotype was 
figured by Walker (1868, pi. 18, fig. 8) and was collected from the Upper Aptian of 
Upware, Cambridgeshire. It has been re-registered as BB. 42904. 

Material and locality. In addition to the holotype, forty-eight specimens 
from Upware registered as B . 25498, B . 25699, B . 25701, B . 25703. In the Davidson 
Collection there are nine specimens from Upware registered as B.6754 and three 
from Brickhill registered as B.6755, all in the British Museum (Natural History). 

Remarks. Often confused with Terebratiila pseudojurensis Leymerie from the 
Middle Neocomian of Marolles, Aube, France, Tamarella vesta nevertheless bears a 
superficial resemblance to this species. It differs fundamentally in its internal 



so Ml': LOWER CRETACEOUS T E R li 15 K A T E L l.O I I ) i: A 

ooooOO 

0-8 0-3 0-2 0-4 ^S^ 0-3 ^P^O-4 



QQOG 



'>-i 




r^r^r~> 




Fig. g. Twenty transverse serial sections through the umbo of Tamarella vesta sp. n. from the 
Upper Aptian of Upware, Cambridgeshire. BM. B. 25700. X2. 



generic characters which can be seen in serial section (Text-fig. 9). It is distinguished 
from T. bonneyi (Keeping) by its more elongate outline, steeper flanks, more produced 
umbo and less prominent concentric growth-lines. It differs from T. juddi and T. 
tamarindus in having flatter or less convex valves and ligate anterior margin. 

A common fossil in the Upper Aptian beds of Upware, T. vesta occurs, though less 
commonly, in beds of equivalent age at Brickhill, Buckinghamshire and at Potton, 
Bedfordshire. 



64 SOME LOWER CRETACEOUS T E R E B R AT E LLC) I I) E A 

Tamarella elliptica (Walker) 

(PI. I, figs. 6a-c) 

i86S Waldheimia miitahilis var. elliptica. Walker : 400, pi. 19, fig. 4. 

1870 Waldheimia mutabilis var. elliptica Walker ; Walker : 562. 

1874 Waldheimia wanklyni var. elliptica Walker ; Davidson : 51, pi. 7, fig.s. 22-25. 

Description. Large oval Tamarella approximately 35 mm. long, 26 mm. wide 
and 15 mm. thick. Both valves are equiconvex and the lateral commissure is 
almost straight. Somewhat resembling T. bonneyi (Keeping) in general outline and 
convexity, T. elliptica lacks the ligation of the anterior margin noted in the former 
species. A large circular foramen truncates a broad, flattened umbo. Sharp beak- 
ridges border a fairly extensive interarea. The shell surface is smooth apart from a 
trace of concentric growth-lines which become more prominent at the margins. 

HoLOTYPE. In the British Museum (Natural History) registered number 67600, 
and figured by Walker (1868, pi. 19, fig. 4) as Waldheimia mutabilis var. elliptica. It 
was collected from the Upper Aptian of Upware. 

Remarks. Walker (1868 : 400) used the name Waldheimia mutabilis for a brachio- 
pod from the Lower Greensand of Upware. No specimen was figured nor was anj^ 
type material indicated. Instead, two forms which Walker called " varieties or sub- 
species " were erected and named W. mutabilis elliptica and W. mutabilis angusta. 
The descriptions of these were accompanied by illustrations (1868, pi. 19, figs. 4, 5) 
of two specimens now in the British Museum (Natural History) and registered as 
67600 and 67601 respective!}'. 

Walker's conception of the species W. mutabilis having been established on the 
description of the two " varieties or sub-species ", is, therefore, indefinable. In any 
case, the name Waldheimia mutabilis had already been used by Oppel (1861) for a 
zeilleriid brachiopod from the Lower Lias of German}-, and it was for this reason that 
Walker asked Davidson (1874 : 51) to correct the name to wankylni in his Cretaceous 
monograph. 

- The varietal name elliptica is here raised to specific rank and referred to Tamarella 
gen. nov. The varietal name angusta has also been raised to specific rank and 
referred to Vectella gen. nov. on p. 53. Both species are common members of collec- 
tions of brachiopods from Upware and are easily recognizable. 

Genus RUGITELA Muir-Wood, 1936 
Rugitela roemeri sp. n. 

(PI. 2, figs, la-c, ^a-c, 6a-c, 8a-c, ga-c ; Text-fig. 10) 

1836 Terehraiitla longa Roemer : 50, pi. 2, fig. 11. 

1839 Terebratiila longa Roemer ; Roemer : 22, pi. 18, fig. 12. 

1840 Terebratiila longa Roemer ; Roemer : 44, No. 50. 

1847 Terebratula faba Sowerby ; d'Orbigny : 77, pi. 506, figs. 8, 9, 11, 12. 

1864 Terebratiila {Waldheimia) faba Sowerby ; Credner : 561, pi. 21, figs. 3, 4, 5. 

1868 Terebratiila longa Roemer ; Quenstedt : 338, pi. 4,6 fig. 99. 

Diagnosis. Rugitela about 28 mm. long, 18 mm. wide and 14 mm. thick. Shell 



SOME LOWER CKETACEOUS T E R E B K AT E LLO I U E A 



05 



biconvex, oval to subtriangular in outline. Umbo short, massive, suberect. Exten- 
sive interarea bordered by sharp, mesothyrid beak-ridges. Foramen large, circular. 
Deltidial plates well exposed, conjunct. Anterior margin rectimarginate to sulcate. 

HoLOTYPE. British Museum (Natural History) no. BB . 42912, from the Neocomian, 
Hilsconglomerat of EUigser Brinke, near Hanover, North German}^ According to 
Judd (1870 : 331) the " Elligser Brinke Schist " has its English equivalent in the 
Acanthodiscus speetonensis Zone at Speeton, Yorkshire and in Lincolnshire. 

Description. This species is characteristically elongate-oval in outline with 
marked growth-lines becoming still more prominent at the margins. Gerontic 
thickening of the anterior and lateral margins is a common feature. Some variants 
show a tendency to broad anterior sulcation of the brachial valve. 

Internal characters. Short, slightly curved, divergent dental lamellae are embedded 
in thick callus and support inwardly curving hinge-teeth. Cardinal process not 
developed. A short septalium develops with long, fused hinge-plates which are 
deflected ventrally forming a small anterior plication. A long median septum exten- 
ding two-thirds the length of the shell supports the hinge-plates. Inner and outer 
socket-ridges are extensive and well marked. Triangular crural bases give rise to 
thick descending branches of the brachial loop with dorsal development and short 
crural processes. Fairly broad transverse band developed from ascending branches 
of the loop. A thickening of the descending branches at a point about one-third the 
distance from the hinge-plates has been noted in mature forms so far sectioned and 
may be due to resorption of the attachment branches to the median septum in the 
dorsal valve. No attachment of brachial loop to septum has been noted in any 
sections of either young or mature forms. 

9 QfQD.QQ 



QQQQQ 



.x^ oK^y oV^ ok^oXLV 

Fig. 10. Eleven transverse serial sections through the umbo of Riigitela roemeri sp. n. 
from the Neocomian of Berklingen, North Germany, showing the characteristic \V -shaped 
hinge-plates and thickened area of the descending branches of the brachial loop. BM. 
B. 35656. X2. 

Remarks. Zieten (1830 : 52) was the first to use the binomen Tercbratula longa 
for a terebratelloid brachiopod which he figured (pi. 39, fig. 7) from the " calcaire 
jurassique " [Upper Jurassic] of Donzdorf, North Germany. Roemer's description 
of Terehratula longa (1836 : 50, pi. 2, fig. 11) from the Neocomian of Hanover was, 
therefore, invalid. 



60 SOiME LOWER CRETACEOUS T E R E 13 R AT ELLO 1 D E A 

D'Orbigny (1847 : 77, pi. 506, figs. 8-12) described and figured a brachiopod from the 
Neocomian of St. Dizier, Haute Marne, France, and assigned it to Terebratitla faba 
Sowerby, a name which J. de C. Sowerby (1836 : 338) had originally used in a des- 
cription of a brachiopod from the Lower Greensand beds between Folkestone and 
Sandgate, Kent. Specimens collected from the Lower Albian beds of Folkestone in 
the Sedgwick Museum (B. 17633-17650), have been matched with others collected 
from the Leymeriella tardcfiircata Zone of Folkestone by Dr. R. Casey and compare 
favourably' in size, general outline and convexity with Terebratula faba as figured by 
J. de C. Sowerby (1836, pi. 14, fig. 10). A duplicate specimen (Brit. Mus (N.H.) no. 
B. 25994) has been sectioned (Text-fig. 12) and is shown to belong to the genus 
Modcstclla Owen (1961 : 573) described from the Lower Albian of Folkestone. 

D'Orbigny's illustrations (1847, pi. 506, figs. 8-12) were probably an idealization of 
eight specimens which were collectively numbered 5159 in the d'Orbigny Collection, 
Laboratoire de Paleontologie, Museum National d'Histore Naturelle, Paris. Unfor- 
tunately only four of these specimens can be traced. One has been identified as a 
specimen of Vedella celtica (Morris) from the Lower Greensand of the Isle of Wight, 
and another (see PI. 2, figs. 8fl-c) is almost identical with the specimen figured by 
Roemer (1839, pl- ^8, fig. 12) from the Neocomian of Hanover. 

Rugitela roemeri occurs rarely in Britain, being found in the upper part of the Claxb}' 
Ironstone, Lincolnshire and in the Speeton Clay of Yorkshire. A fine, well preserved 
specimen (PL 2, figs, la-c) from the Claxby Ironstone of Nettleton, Lincolnshire, 
collected by Mr. Peter Rawson of Hull University, is the only example known from 
these beds which compares exactly with Roemer's original illustration (1836, pi. 2, 
fig. 11). The dimensions of this specimen are : 40 mm. long, 20 mm. wide and 16 
mm. thick. 

A form described here as Rugitela rugosa sp. n. resembles R. roemeri but is shorter, 
more gibbous and oval in outline with no marked sulcation of the anterior portion of 
the brachial valve. It is found in the same beds of the Claxby Ironstone and is 
probably present in the Speeton Clay, although specimens collected from beds within 
the range C6 to C3 of Swinnerton (1936-55) and which may belong to this species are 
too crushed for accurate determination. R. rugosa also occurs in the Hanterivian at 
Salins-les-Bains in the French Jura Mountains. A well preserved example is figured 
here (PI. 2, figs, ^a-d) for comparison. 

The reference of this species to the genus Rugitela has been made after careful 
comparison with transverse serial sections of the type-species R. bullata Muir-Wood 
and extends the range of the genus from ? Inferior Oolite to the Lower Cretaceous. 
It is possible that the final range may include species from the Inferior Oolite at 
present under review by Dr. H. M. Muir-^^"ood. 

Material and locality. Well preserved material from the German Neocomian 
is rare but there are, in addition to the holotype metioned above, nine other specimens 
in the Davidson Collection, Brit. Mus. (Nat. Hist.) B.6759, BB.41911, from the 
" Elligser Brinke Schist " of Delligsen near Hanover. 

A specimen from the Neocomian of St. Dizier in the Haute-Marne is also in the 
Davidson Collection (B.6736). It is accompanied by the note in Davidson hand- 
writing which reads : "... given to me b}' d'Orbigny . . . from the d'Orbigny 
Coll ..." 



SOME L O \N' K R C R E '1' A C E O U S 1' li R li B R A T E L L O 1 D E A 



67 



Seven specimens, British Museum (Natural History) (BB. 42925-31), from the 
Acanthodiscus speetonensis Zone of the Speeton Clay, Yorkshire. Although crushed 
they compare favourably with the North German forms. 

Rugitela rugosa sp. n. 

(PI. 2, figs. 2a-c, ^(i-c, ^a-c ; Text-fig. 11) 

1874 Waldheimia f aba d'Orhigny ; Davidson : pi. 6, figs. 12-14. 

Diagnosis. Biconvex Rugitela about 29 mm. long, 17 mm. wide and 17 mm. thick. 
Oval to sub-circular in outline. Shell surface covered with pronounced growth- 
lines. Anterior and lateral margins thickened. Umbo short, slightly incurved. 
Interarea extensive. Deltidial plates conjunct. Beak-ridges sharp. Anterior 
commissure, rectimarginate. 

HoLOTVPE. Davidson Collection, British Museum (Natural History) no. B.6734 
from the Claxby Ironstone (Lower Hauterivian) of Acre House, Tealby, Lincolnshire. 

Material. In addition to the holotype, four specimens from the Claxby Iron- 



000000 







• t 




/( 



• • 



0-4 




0-5 



Fig. II. A scries of sixteen transverse sections through the umbonal part of Rugitela rugosa 
sp. n. from the Ncocomiau, Claxby Ironstone, Acre House, Tcalb\-, Lincolnshire. 
BM.B. 50359. X2. 



68 



SOME LOWER CRETACEOUS TE RE B R A TELLO I D E A 



stone, British Museum (Natural History) nos. B . 6734, B . 50358-59, B . 50324 and two 
well preserved specimens from Salins-les-Bains, nos. 33984 and BB. 42913. There 
are two specimens from the Claxby Ironstone in the Sedgwick Museum (B .11400 and 
B. 12278). 

Description. In general outhne this is a shorter, more gibbous species than 

o oOO 

0-4 0-2 >y^o-4 ^ti^O-S 
0-4 0-6 ^*^0-3 





Fig. 12. Eleven transverse serial sections through the umbo of Alodestella Jala (J. de C. Sowerby) 
from the f.ower Albian, Folkestone, Kent. BM. B. 25994. x 4. 



SOME LOWER CRETACEOUS TERE B R ATE LLO I D E A 69 

Ritgitela roemeri. It has more acutely convex vah'cs with an absence of any marked 
sulcation of the brachial valve, although some mature forms exhibit a faint depression 
in this part of the shell. The surface of the valves is ornamented by numerous con- 
centric growth-lines which give it a rugose appearance. Marginal thickening of the 
valves is a common feature in more mature forms. The umbo is less produced and 
more incurved than in R. roemeri. Well defined, sharp beak-ridges border an 
extensive inter area. 

Locality and horizon. This species enjoys the same lateral range as Rugitela 
roemeri, occurring in the Lower Hauterivian at Salins-les-Bains in the French Jura 
Mountains ; a similar horizon at Brunswick, North Germany, and in the Claxby 
Ironstone of Lincolnshire, as well as the Acanthodiscits speetonensis Zone of the 
Speeton Clay of Yorkshire. 

Rugitela hippopus (Roemer) 

(PL 3, figs, la-c, 20,-0, 3a-c, ^a-c ; Text-fig. 13) 

1840 Terebratida hippopus Roemer : 114, pi. 16, fig. 28. 

1842 Terebratida hippopus Roemer ; Geinitz : 87. 

1864 Terebratida [W aldheimia) hippopus Roemer ; Credner : 565, pi. 21, figs. 17, 18, 19. 

1866 Terebratida hippopus Roemer ; Schloenbach : 33. 

1868 Terebratida ? deflitxa Schloenbach : 31, pi. 2, figs. 10-12. 

1868 Terebratida hippopus Roemer ; Judd : 245. 

1874 Waldheimia hippopus ? Roemer var. tilbyensis Davidson : 53, pi. 6, figs. 10, 11. 

1884 Terebratida [Waldheimia) hippopus Roemer; Weerth : 62, pi. 11, figs. 5, 6. 

Description. This well established species is characterized by its almost circular 
outline, sulcate brachial valve, slightly incurved beak and acutely carinate pedicle 
valve. It is 12 mm. long, 10 mm. wide and 8 mm. thick. Because of the similarity 
of external form it has often been assigned to the Liassic genus. 

Aulacothyris. It differs from this genus, however, in having a less elongate outline, 
smaller dimensions, larger foramen and longer brachial loop. 

Internal characters. From the transverse serial section of the species (Text-fig. 13) 
it is possible to compare the internal characters of R. hippopus with sections of species 
of Rugitela figured by Muir-Wood (1936 : 124, 128, 131). While agreeing basically 
with sections of the type-species Rugitela bullata from the Fuller's Earth Rock of 
Somerset, it appears to have closer morphological affinities with R. emarginata from 
the same horizon and locality. 

Neotype. Dr. R. G. Thurrell of H.M. Geological Survey has kindly given me 
permission to publish some of the notes he prepared in a revision of this species which 
he included in his thesis for the degree of Ph.D of London University, 1957. The 
following is a quotation from p. 320 of his thesis : 

" Owing to war damage, the Roemer Collection in Hildesheim Museum was partly 
destroyed. Dr. Friedrich Smid of the German Geological Survey has undertaken to 
search for the type material, but nothing has so far come to light." 

In spite of renewed enquiry no specimens have been forthcoming. 

A specimen in the British Museum (Natural History) no. 32313 from the Hilscon- 
glomerat of Berklingen, North Germany (PI. 3, figs. 2a-c) appears to be the only 



70 



SOME LOWER CRETACEOUS TE RE BR AT ELLO I D E A 

o o o 

^'^O-S ^^0-2 ^*->^0-4 ^^•-^^0-3 ^%^^o 

O. O. Q. O 



Fig. 13. A series of eleven transverse serial sections through the umbo of Riigitela hippopus 
(Roemer) from the Neocomian, Claxby Ironstone, Acre House, Tealby, Lincolnshire. 
BM.BB. 42934. X2. 

available specimen from the type locality, and is here proposed as neotype. It 
departs from the original illustration (Roemer 1840, pi. 16, fig. 28) in having less 
clearly defined beak-ridges and less extensive interarea but, after examination of 
some 150 specimens from both English and North German localities, these features 
appear to vary within the species. 

Material and locality. Numerous specimens from the Claxby Ironstone from 
Acre House, Tealby and from Nettleton, Lincolnshire. Ten specimens from the 
Neocomian of Saltzgitter, Hanover (B.6748, BB. 42921, BB. 42932), a single speci- 
men from the Speeton Clay of Yorkshire (BB. 42923), and one specimen from 
MaroUes, Aube, France (B.6742), all in the British Museum (Natural History). 

Remarks. Differing only in size from the North German form, the British species, 
originally described as Waldheimia hippopus var. tilbyensis by Davidson (1874), 
attains an average length of 15 mm., a width of 13 mm., and a thickness of 10 mm. 

The specimen from the Lower Hauterivian of Marolles, France, differs slightly in 
beak characters. It has less prominent beak-ridges and less extensive interarea. 

conclusion 

Owing to the lack of recent exposures of the Lower Greensand at Upware, Brickhill 
and Potton, it is impossible to draw any real conclusions about the distribution of 
species. All information regarding these localities has been obtained from research 
on museum collections and the results recorded here. 

Critical examination of transverse serial sections of species of Tamarella gen. nov. 
has revealed that they have many characters in common with those of Rugitela. 
For example, the divergent dental lamellae, broad, shallow septalium, anteriorly 



SOiME LOWER CRETACEOUS T E R E 13 R A TE LLO 1 D E A 71 

arched and fused hinge-plates, triangular crural bases and thickened portion of the 
descending branches of the brachial loop. The dorsal median septum of Ritgitela 
is, however, more strongly developed and considerably longer than that of Taniarella. 
Prozorovskaya (1962 : in) described Gusarella from the Upper Jurassic of Turk- 
menistan as a subgenus of Zeilleria. Serial sections of the type species Z. {Gusarella) 
gusarensis have much in common with those of Taniarella particularly T. bonneyi 
(Text-fig. 7). The hinge-plates show the anterior arching or plication, and the crural 
bases, brachial loop and general outline are similar to those of Taniarella. In 
Gusarella, however, the dorsal median septum appears to be absent. 

ACKNOWLEDGMENTS 

I am grateful to Dr. H. M. Muir-Wood for her kindness and generosity in many ways 
and help with the manuscript and also for her continued encouragement. 

I am also grateful to Dr. W. T. Dean for many helpful suggestions and for some of 
the photographs. My thanks are also due to Mr. John Ferguson for photographic 
assistance and helpful discussion ; to Mr. Peter Rawson of the Dept. of Geology, 
Hull University ; to Dr. Colin Forbes, Sedgwick Museum, Cambridge and to Dr. R. G. 
Thurrell of H.M. Geological Survey. 

REFERENCES 

Casey, R. 1961. The stratigraphical palaeontology of the Lower Greensand. Palaeontology, 

London, 3 : 487-621, pis. 77-84. 
Cooper, G. A. 1955. New Cretaceous Brachiopoda from Arizona. Smithson. Misc. Coll., 

Washington, 131, 4 : 1-18, pis. 1-4. 
Credner, H. 1864. Die Brachiopoden der Hilsbildung im nordwestlichen Dcutschland. Z. 

dtsch. geol. Ges., Berlin, 16 : 542-572, pis. 18-21. 
Davidson, T. 1852-55. A Monograph of British Cretaceous Brachiopoda, 1. 117 pp., 12 pis. 

[Mon. Palaeont. Soc., London.] 

1874. A Monograph of the British Fossil Brachiopoda, 4, i. Supplement to the Recent, 

Tertiary, and Cretaceous Species. 72 pp., 8 pis. [Mon. Palaeont. Soc, London.] 

Davidson, T. & Morris, J. 1847. Descriptions of some species of Brachiopoda. Ann. Mag. 

Nat. Hist., London (i) 20 : 250-257. 
FiTTON, J. 1836. Observations on some of the strata between the Chalk and Oxford Oolite 

in south-east of England. Trans. Geol. Soc. Lond. (2) 4 : 335-349, pis. 11-23. 
Geinitz, H. B. 1842. Ueber Versteinerungen Altenburg und Ronneburg. Mitth. Osterl. 

Altenburg, 6 : 86-99. 
JUDD, J. W. 1868. On the Speeton Clay. Quart. J. Geol. Soc. Lond., 24 : 218-250. 

1870. Additional observations on the Neocoraian strata of Yorkshire and Lincolnshire, 

with notes on their relations to the beds of the same age throughout Northern Europe. 
Quart. J . Geol. Soc. Lond., 26 : 326-348. 

Keeping, W. 1883. The fossils and palaeontological affinities of the Neocomian deposits of 
Upware and Brickhill. Sedgwick Prize Essay for 1879. xi + 167 pp., 8 pis. 

Lankester, E. R. 1863. On certain Cretaceous brachiopoda. Geo/o^w/, London, 6 : 414-415. 

Leymerxe, A. 1841-42. Memoire sur le terrain Cretac6 du Departement de I'Aube, contenant 
des considerations gen^rales sur le terrain N^ocomien. Mem. Soc. geol. Fr., Paris (i) 4 
(1841) : 291-364, pis. 16-18, 5 (1842) : 1-34, pis. 1-18. 

Meyer, C. J. A. 1864. Notes on the Brachiopoda from the Pebble-bed of the Lower Green- 
sand of Surrey ; with descriptions of the new species, and remarks on the correlation of the 
Greensand Beds of Kent, Surrey and Berks., and of the Farringdon Sponge-gravel, and the 
Tourtia of Belgium. Geol. Mag., London (i) 1 : 249-257, pis. 11-12. 



72 SOME LOWER CRETACEOUS TEREB R ATELLO 1 DE A 

Meyer, C. J. A. 1868. Notes on Cretaceous Brachiopoda and on the development of the loop 

and septum in Terebratella. Geol. Mag., London (i) 5: 268-272. 
MiDDLEMiss, F. 1959. English Aptian Terebratulidae. Palaeontology, London, 2 : 9.^-142, 

pis. 15-18. 
Morris, J. 1843. A Catalogue of British Fossils, comprising all the genera and species hitherto 

described, with references to their geological distribution and to the localities in which they have 

been found, x + 222 pp. London. 
1854. A Catalogue of British Fossils, comprising all the genera and species hitherto described, 

with references to their geological distribution and to the localities in which they have been found. 

2nd ed. vii + 372 pp. London. 
MuiR-WooD, H. M., 1934. On the internal structure of some Mesozoic Brachiopoda. Philos. 

Trans., London (B) 223 : 511-567, pis. i, 2. 
1936. Brachiopoda of the British Great Oolite Series. I. The Brachiopoda of the Fuller's 

Earth, ii + 144 pp., 5 pis. London. 
Oppel, a. 1861. Ueber die Brachiopoden des untern Lias. Z. dtsch. geol. Ges., Berlin, 13 : 

529-550. pis. 1-4- 
Orbigny, A. D'. 1848-51. Terrains cretaces. Paleojitologie frajicaise, 4 : 1-390, pis. 490-599. 

Paris. 
Owen, E. F. 1961. In Casey, R. The stratigraphical palaeontology of the Lower Greensand. 

Palaeontology, London, 3 : 487-621, pis. 77-84. 
1963. The brachiopod genus Modestella in the Lower Greensand of Great Britain. Ann. 

Mag. Nat. Hist., London {13) 6 : 199-203, pi. 10. 
Prozorovskaya, E. L. 1962. Some new brachiopods found in the Upper Jurassic deposits of 

Western Turkmenistan. ]'est. Leningr. Univ. Leningrad, 12 (Geol. & Geogr.) 2 : 108-114. 
Quenstedt, F. A. 1868-71. Petrefaktenkunde Deutschlands. 11 Brachiopoden. iv + 748 pp., 

25 pis. Tubingen. 
RoEMER, F. A. 1836. Die V'ersteincrungen des Norddeutschen Oolithen-Gebirges. 218 pp., 

16 pis. Hannover. 

1839. Die Versteinerungen des Norddeutschen Oolithen-Gebirges . 59 pp., 4 pis. Hannover. 

1840. Die Versteinerungen des Norddeutschen Kreidegebirge. iv + 145 PP-. 16 pis. 

Hannover. 

ScHLOENBACH, U. 1866. Ueber die Brachiopoden aus dem unteren Gault (Aptien) von Ahaus 

in Westphalen. Z. dtsch. geol. Ges., Berlin, 18 : 364-376. 
ScHLOENBACH, U. 1868. Ubcr die norddeutschen Galeriten-Schichtcn und ihre Brachiopod 

Fauna. S.B. K. Akad. Wiss. Wien, 57 : 181-224, P^^- i-3- 
SowERBY, J. de C. 1836. see Fitton, J. 
SwiNNERTON, H. H. 1936-55. A Monograph of British Cretaceous Bclcmnitcs. 86 pp., i8 pis. 

[Mon. Palaeont. Soc, London.] 
Walker, J. F. 1867. On some new Terebratulidae from Upwarc. Geol. Mag., London (i) 

4 : 454-456, pi. 19- 

1868. On the species of Brachiopoda which occur in the Lower Greensand at Upware. 

Geol. Mag., London (i) 5 : 339-406, pis. 18-19. 

1870. On secondary species of Brachiopoda. Geol. Mag., London (i) 7 : 560-564, pi. i. 

Weerth, O. 1884. Die Fauna des Neocomsandsteins im Teutoburger Walde. Paldont. Abh., 

Berlin, 2 : 1-77, pis. i-ii. 
Zieten, C. H. V. 1830-33. Die Versteinerungen Wurttenibergs . 1-16, pis. 1-12 (1830) ; 

17-32 pis. 13-24 (1831) ; 33-64, pis. 25-48 (1832) ; 65-102, pis. 49-72 (1833). Stuttgart. 



PLATE I 
a. Dorsal view. b. Lateral view. c. Anterior view. 

Figs. la, b, c. Vectella woodwardi (Walker), L^ppcr Aptian of Upware, Cambridgeshire. 
BM.BB. 42922. 

Figs. 2a, b, c. TamareUa tamarindiis (J. de C. Sowerby), Upper Aptian, L'pware, Cambridge- 
shire. BM.BB. 42905. 

Figs. 3a, b, c. TamareUa bonnevi (Keeping), Upper Aptian, Brickhill, Buckinghamshire. 
BM.BB. 42935. 

Figs. 4a, b, c. Vectella celtica (Morris), L^pper Aptian, Shanklin, Isle of \Yight. B]\L BB. 42915. 
Lectotj'^pe. 

Figs. 5a, b, c. Vectella angitsta (Walker), L'pper Aptian, LTpwarc, Cambridgeshire. B^L no. 
67601. Holotype. 

Figs. 6a, b, c. TamareUa elliptica (Walker), LTpper Aptian, L^pware, Cambridgeshire. B]NL 
no. 67600. Holotype. 

Fig. 7. Vectella morrisi (Meyer), Upper Aptian, Bargate Pebble-bed, Compton, Surrey. 
Internal of brachial valve showing the broad, deep hinge-trough and extensive dorsal median 
septum. BM. BB. 42918. 

Figs. 8a, b, c. TamareUa tamarindiis (J. de C. Sowerby). Dissected specimen showing rela- 
tively shallow hinge-trough in brachial valve 8a, with anterior plication and 8b. pedicle valve of 
the same specimen. BM BB. 42920. 8c. Dissected specimen of a brachial valve of a specimen 
from the Upper Aptian of Faringdon. BM. BB. 42919, showing cardinalia and remains of brachial 
loop with thickened portion of descending branch. 

Figs, ga, b, c. TamareUa vesta sp. n., Upper Aptian, Upware, Cambridgeshire. B^L 
BB. 42904. Holotype. 

Figs. loa, b, c. TamareUa tamarindiis (J. de C. Sowerbv), L'pper Aptian, Shanklin, Isle of 
Wight. BM.BB. 42906. 

Figs, iia, b, c. TamareUa jiiddi (Walker), L'pper Aptian, L'pware, Cambridgeshire. B^I. 
BB. 42909. Lectotype. 



BitlLBM. (N.H.) Geol. 11,2 
3^ 



PLATE I 




PLATE 2 
a. Dorsal view. b. Lateral view. c. Anterior view. 

Figs. la, b, c. Rugitela voemeri sp. n. from the Neocomian (Hauterivian), Nettleton, Lincoln- 
shire. Hull University Collection H.U.C/Rn. 317. 

Figs. 2a, b, c. Rugitela rugosa sp. n., Neocomian, Claxby Ironstone, Acre House, Tealby, 
Lincolnshire. BM.B. 50324. 

Figs. 3a, b, c. Rugitela roemeri sp. n., Neocomian, St. Dizier, Haute-Mame, France. BM. 
B.6736. 

Figs. 4a, b, c. Rugitela nigosa sp. n., Neocomian, Claxby Ironstone, Acre House, Tealby, 
Lincolnshire. BM. B.6734. Holotype. 

Figs. 5a, b, c. Rugitela rugosa sp. n., Neocomian, Salins-les-Bains, Jura Mountains, France. 
BM.BB. 42913. 

Fig. 5d. Enlarged portion of above specimen showing incurved beak, sharp beak-ridges and 
wide interarea. 

Figs. 6a, b, c. Rugitela roemeri sp. n., Neocomian, Elligser Brinke, Hanover, North Ger- 
many. BM. BB. 42912. Holotype. 

Figs. 7a, b, c. Modestella faba (J. de C. Sowerby), Lower Albian, Folkestone, Kent. SM. 
B. 17642. 

Figs. 8a, b, c. Rugitella roemeri sp. n., Neocomian, St. Dizier, Haute-Mame, France. 
d'Orbigny Collection no. 5159. 

Figs, ga, b, c. Rugitela roemeri sp. n., Neocomian, Elligser Brinke, Hanover, North Ger- 
many. BM.BB. 42911. 



Bull. B.M. [N.H.) Geol. 11, 2 



PLATE 2 




PLATE 3 
a. Dorsal view. b. lateral view. c. Anterior view. 

Figs. la, b, c. Riigitela hippopus (Roemer), Neocomian, Speeton Clay, Speeton, Yorkshire. 
P. Rawson Coll. BM. BB . 42923. x 2. 

Figs. 2a, b, c. Rngitela hippopus (Roemer), Neocomian, Berklingen, near Hanover, North 
Germany. BM. no. 32313. Neotype. X2. 

Figs. 3a, b, c. Rngitela hippopus (Roemer), Neocomian, Saltzgitter, North Germany. BIM 
BB. 42921. X2. 

Figs. 4a, b, c. Rngitela hippopus (Roemer), Neocomian, Saltzgitter, North Germany. BM 
BB. 42932. X2. 

Figs. 5a, b, c. Tamarella tamarindiis (J. de C. Sowerby), Upper Aptian, Isle of Wight, BM 
BB. 42907. Neotype. X2. 

Figs. 6a, b, c. Tamarella tamarindus (J. de C. Sowerby), Upper Aptian, Faringdon, Berkshire 
BM. B.6723. 

Figs. 7a, b, c. Vectella moyyisi (Meyer), Upper Aptian, Brickhill, Buckinghamshire. BM 
BB. 42916. X2. 

Figs. 8a, b, c. Vectella morrisi (Meyer), Upper Aptian, Godalming, Surrey. BM. BB.42917 

X2. 

Figs. 9a, b, c. Vectella morrisi (Meyer), Upper Aptian, Shanklin, Isle of Wight, BM. BB . 42914 
Fig. 9d. Vectella morrisi (Meyer), Upper Aptian, Shanklin, Isle of Wight. SM. B. 14770 
Lectotype. 



Bull.BM. {N.H.) Geol. 11,2 



PLATE 3 







PRINTED IN GREAT BRITAIN 
BY ADLARD & SON LIMITED 
BARTHOLOMEW PRESS, DORKING 



4 ■ N 

MIDDLE JURASSIC OSTRACODA v%jj^< 
FROM THE GREY LIMESTONE 
SERIES, YORKSHIRE 



R. H. BATE 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. ii No. 3 

LONDON: 1965 



MIDDLE JURASSIC OSTRACODA FROM THE /rTl^ 
GREY LIMESTONE SERIES, YORKSHIRE '^ 



BY 

R. H. BATE 



\^i 



Pp. 73-133 ; 21 Plates ; 24 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

GEOLOGY Vol. 11 No. 3 

LONDON: 1965 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 3 of the Geological 
(Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 



Trustees of the British Museum (Natural History) 1965 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 8 December, 1965 Price £4 



MIDDLE JURASSIC OSTRACODA FROM THE 
GREY LIMESTONE SERIES, YORKSHIRE 

By RAYMOND HOLMES BATE 



CONTENTS 

I Introduction and acknowledgements 

II Stratigraphical sections . 

III Stratigraphy . 

IV Conclusions 
V Systematic descriptions . 

Subclass Ostracoda Latreille 
Order Podocopida Miiller 
Suborder Podocopina Sars . 

Superfamily Cypridacea Baird. 
Family Paracyprididae Sars 
Genus Paracypris Sars . 

Paracypris bajociana Bate 
Superfamily Cytheracea Baird . 
Family Bythocytheridae Sars 
Genus Monoceratina Roth 

Monoceratina scarboroughensis sp. nov. 
Family Progonocytheridae Sylvester-Bradley. 
Subfamily Progonocytherinae Sylvester-Bradley 
Genus Caytonidea nov. .... 

Caytonidea faveolata sp. nov. 
Genus Cloughtonella nov. 

Cloughtonella rugosa sp. nov. . 
Genus Fuhrbergiella Brand & Malz 

Subgenus Praefuhrbergiella Brand & Malz 

Fuhrbergiella [Praefuhrbergiella) horrida 
horrida Brand & Malz 
Genus Glyptocythere Brand & Malz 
Glyptocythere costata sp. nov. 
Glyptocythere polita sp. nov. 
Glyptocythere scitula sp. nov. 
Genus Malzia nov. 

Malzia bicarinata sp. nov. . 
Malzia unicarinata sp. nov. 
Genus Progonocythere Sylvester-Bradley 
Progonocythere acuminata sp. nov. 
Progonocythere yonsnabensis sp. nov. 
Subfamily Pleurocytherinae Mandelstam . 
Genus Pleurocythere Triebel . 
Pleurocythere sp. 



page 

76 
80 
92 

97 
98 
98 
98 
98 
98 
98 
98 
98 
99 
99 
99 
99 
100 

ICO 

100 
100 

lOI 

102 
104 
104 

104 
106 
106 
107 
108 
no 
III 

"3 
114 
114 
116 
117 
117 
117 



GEOL. II, 3 



76 



R. H. BATE 



VI 



Family Cytherideidae Sars . 

Subfamily Cytherideinae Sars . 
Genus Vernoniella Oertli 

Vernoniella bajociana sp. nov. 
Vernoniella ? caytonensis sp. nov 
Genus Ljubimovella Malz 

Ljiibimovella piriformis Malz 
Family Schulerideidae Mandelstam 
Subfamily Schulerideinae Mandelstam. 
Genus Mesocytheridea nov. 

Mesocytheridea howardianensis sp. nov 
Genus Praeschuleridea Bate . 

Praeschuleridea subtrigona (Jones & Sherbom) 

Praeschuleridea subtrigona intermedia subsp 

nov. ..... 

Family Cytheruridae Miiller .... 

Genus Eocytheropteron Alexander . 

Eocytheropteron ? sp. . 
Genus Paracytheridea Miiller . 

Paracytheridea ? caytonensis sp. nov. 
Family Protocytheridae Ljubimova 
Subfamily Kirtonellinae Bate . 

Genus Southcavea Bate .... 

Southcavea microcellulosa sp. nov. 
Genus Systenocythere Bate 

Systenocythere ovata sp. nov. 
References ........ 



page 

117 
117 
117 
118 
119 
120 
120 
121 
121 
121 
122 
124 
124 

124 
126 
126 
126 
127 
127 
128 
128 
128 
128 
130 
130 
132 



SYNOPSIS 

The Middle Jurassic (Bajocian) Grey Limestone Series of Yorkshire is shown to have been 
deposited in a broad embayment, transgressing deltaic sediments of Bajocian age, and is here 
divided into two ostracod faunal zones : a lower zone of Glyptocythere polita and an upper zone of 
G. scitula, the type section being at Hundale Point. The zone of G. polita is present only 
towards the centre of the basin and represents the first phase of the marine transgression. The 
facies changes within the Series are produced by intermittently rising sea-level and the repeated 
southward extension of the northern delta. Shore-line sediments are also identified. Fourteen 
stratigraphical sections are described in detail. The period of marine deposition is considered to 
have taken place approximately during sauzei to blagdeni times and is thus in part at least 
equivalent to the Coronaten-Schichten of N.W. Germany. The ostracod fauna of the Grey 
Limestone Series is described for the first time. Four genera : Caytonidea, Cloughtonella, Malzia 
and Mesocytheridea are new. Sixteen new species and one new subspecies are also described. 
The palaeoecology of the ostracod fauna is briefly mentioned. 



I INTRODUCTION AND ACKNOWLEDGEMENTS 

In north-eastern England, the uppermost marine horizon of Bajocian age consists 
of a well developed succession of calcareous shales and sandy limestones known as the 
Grey Limestone Series or Scarborough Beds. The maximum thickness of this 
sequence occurs in the neighbourhood of Cloughton Wyke (62 feet) and Ravenscar 
(62 feet), the beds thinning to the north, south and west where they eventually pass 



MIDDLE JURASSIC OSTRACODA 



77 



NORTH 
SEA 




Fig. I. Outcrop of Middle Jurassic Strata in N.E. England, with the localities and sections 

(1-14) mentioned in the text. 



78 



R. H. BATE 



into arenaceous shore-line deposits. The open sea during this period lay to the east, 
occupying the site of the present day North Sea, whilst a delta lay to the north. 
Sediments equivalent in age to the Grey Limestone Series were also deposited in 
north-western Germany. 

The dating of the Grey Limestone Series is difficult owing to the almost complete 
absence of ammonites : those which have been recorded (Fox-Strangways 1892 : 231 ; 
Buckman igii : 205-208 ; and Hemingway 1951 : 119)^ indicate that part of the 
sequence at least belongs to the Teloceras blagdeni Subzone. As the ammonite 
zones are well known in north-western Germany, comparison of the ostracod faunas 
of the two regions should eventually offer a solution to this problem. At the moment, 
however, only two ostracods, present in the Grey Limestone Series have also been, 
recorded from Germany ; these are Ljubimovella piriformis Malz (1961 : 165, pi. 2, 
figs. 15-25) and Fuhrhergiella {Praefiihrhergiella) horrida horrida Brand & Malz 
(1962 : 19, pi. 4, figs. 33-37 ; pi. 5, fig. 46). 

Ljubimovella piriformis has a range in Germany of sauzei to blagdeni Zones, with 
the type horizon being in the saiizei Zone. In Yorkshire this ostracod is restricted to 
the zone of Glyptocythere scitula, the upper zone from which the ammonites recorded 
from the Grey Limestone Series were most probably obtained. The ostracod Fuhr- 
hergiella {Praefuhrbergiella) horrida horrida is recorded from the upper part of the 
Coronaten-Schichten {romani to blagdeni Zones) in Germany, whilst in Yorkshire it is 



Table i 
Generalised table of the Middle Jurassic stata in Yorkshire, north of Market Weighton 



Strata 


Stage 


CORNBRASH (UppER) 


Callovian 


Upper Deltaic Series 


Bathonian 






Grey Limestone Series 




Middle Deltaic Series (Upper) 




MiLLEPORE Series 




Middle Deltaic Series (Lower) 


Bajocian 


Eller Beck Bed /Hydraulic Limestone 




Lower Deltaic Series 




Dogger 




Lias 


Toarcian 



^ The map reference for Monk's Walk Wood, Sneaton has been given with a misprint, and should read 
NZ/896086 (personal communication. Prof. Sylvester- Bradley). 



) 


V ] 




~^ 




UPPER DELTAIC 






3 



A 



I'm. a. Scctiuns 1-6 ucrosa llw Yorksliiic Iteiu. 



MIDDLE JURASSIC OSTRACODA 79 

largely restricted to the G. scitiila zone, being found at only one locality in the G. 
polita Zone. 

The Grey Limestone Series represents a period of deposition which appears to 
approximate with the Coronaten-Schichten of Germany although there is insufficient 
evidence to say how many of the German ammonite zones are represented in the 
Yorkshire sediments. It should be possible, however, to correlate the Grey Lime- 
stone Series with the German succession more precisely at a future date when the 
ostracod faunas are more fully known. 

Fox-Strangways (1892 : 223) recorded the presence of lamelUbranch casts within a 
15 ft. bed of sandstone at Ravenscar. This is bed 17 of Section 5 (Text-fig. 2), which is 
the lower part of the Moor Grit (basal Upper Deltaic Series) . The period of marine 
deposition was thus brought to a close by the infilling of the Yorkshire basin with 
deltaic sediments. The next marine transgression did not occur until Callovian times, 
it is, therefore, evident that the end of the Bajocian and the whole of the Bathonian 
is represented in Yorkshire by deltaic sediments of the Upper Deltaic Series. 

The localities listed below are those at which the Grey Limestone Series has been 
examined at outcrop, the numbers corresponding with those in Text-fig. i. Map 
references refer to the one-inch Ordnance Survey map of Great Britain (seventh 
series) . 

Coastal Exposures 

1. Gristhorpe Bay, cliff section and foreshore, map reference TA/085842. A 

complete section, maximum thickness 15 ft. 6 in. 

2. Cayton Bay, cliff section, map reference TA/082843. A complete section, 

maximum thickness 10 ft. 7 in. 

3. White Nab, Scarborough, foreshore section, map reference TA/058865. Base 

below Low Water Mark, seen to 25 ft. 6 in. 

4. Hundale Point, Cloughton Wyke, cliff and foreshore section, map reference 

TA/024949 to TA/027943. A complete section, maximum thickness 62 ft. 
4 in. 

5. Ravenscar, cliff section, map reference NZ/988012. A complete section, maxi- 

mum thickness 62 ft. 4 in. 

Inland Exposures 

6. Hawsker, stream section at Hawsker Bottoms, map reference NZ/937079. Top 

of section not seen, 22 ft. 5 in. 

7. Bloody Beck, stream section, map reference SE/945981 to SE/947982. A 

complete section, 29 ft. 4 in. 

8. May Beck, stream section, map reference NZ/890015 to NZ/891019. Isolated 

exposures, base not seen, 14 ft. 9 in. 
g. Ramsdale Beck, stream section NZ/925034. Upper part of Series only exposed, 
seen to 9 ft. 6 in. 



8o R. H. BATE 

10. EUer Beck, just downstream of road bridge, map reference SE/856984. Upper 

part of Series only, seen to 6 ft. 5 in. 

11. Harland Beck, stream section, map reference SE/684914 to SE/686913. In- 

complete section seen to 14 ft. 4 in. 

12. Bogmire Gill, stream section, map reference SE/608905 to SE/609909. Almost a 

complete section, seen to 40 ft. 3 in. 

13. Yearsley Moor, map reference SE/579754. A small quarry by the side of the 

Yearsley- Ample forth road. Faima indicates position of beds high up in the 
Grey Limestone Series, seen to 8 ft. 3 in. 

14. Stonecliff Wood, Crambeck, map reference SE/744676, a small guUy exposing 

12 ft. 2 in. of sediment. Map reference SE/743676, section higher up the bank, 
above the York-Scarborough railway-line, exposing 5 ft. 8 in. of limestone. 
Map reference SE/740675 a 9 inch bed of limestone is exposed. All strata high 
up in the Grey Limestone Series. 

Morphological terms used in the text are those introduced in vol. Q of the American 
Invertebrate Paleontology Treatise (Moore 1961), Sylvester-Bradley (1956) 
and Bate (1963). 

The work embodied in this paper was largely conducted in the Geology Depart- 
ments of Sheffield University and Leicester University. My sincere thanks go to 
Professors L. R. Moore and P. C. Sylvester-Bradley respectively, for the use of their 
departmental facilities. Grateful thanks are also due to the Department of Industrial 
and Scientific Research for a grant to finance this work. Dr. H. Malz, Senckenberg 
Museum, Frankfurt -am-Main and Dr. H. J. Oertli, S.N.P.A., Centre de Recherches, 
Pau (Basses-Pyr) also assisted with the loan of type and/or comparative material, 
for which they are gratefully acknowledged. 

II STRATIGRAPHICAL SECTIONS 

Section No. i. Gristhorpe Bay (Text-fig. 2), exposure in south face 
of Yons Nab headland and on the foreshore at low tide. The 
section is complete and exposes 15 ft. 6 in. of sediment (G.L.S.). 

Upper Deltaic Series 

ft. in. 
10. Yellow-grey sandstone (Moor Grit) with coaly lenses and grey sul- 
phurous shale . . . . . . . . . 30-40 o 

Grey Limestone Series 

9. Black, sulphur stained shale ....... 4 o 

8. Dark grey, ironstained shale ....... 4 9 

7. Mudstone .......... o 9 

6. Black shale with indet. ostracods ...... i 4 



UPPER DELTAIC 
SERIES 



LIMESTONE 



SERIES 



SERIES 








d 

-^ 



■ Bt'Wi 



I I 

I I 

I I 






l'"!(i. 3. Sfclioiis 7 luul ia-14. 



4 


10 


o 


5 


7 






MIDDLE JURASSIC OSTRACODA 8r 

Jt. in. 

5. Fossiliferous mudstone with Glyptocythere scitula ; Vernoniella 

bajociana and Monoceratinascarboroughensis .... 6 

4. Black fossiliferous shale with a compact limy band at base. Lamil- 

libranch Gervillia scarburgensis and Belemnites common. Glypto- 
cythere scitula ; Praeschuleridea subtrigona intermedia ; Vernoniel- 
la bajociana ; Vernoniella ? caytonensis ; Systenocythere ovata ; 
Paracypris bajociana ; Caytonidea faveolata ; Fuhrbergiella 
[Praefurbergiella) horrida horrida and Pleurocythere sp. . -4 6 

Middle Deltaic Series 

3. Yellow sandstone, flaggy in part with carbonaceous bands 

2. Black Shale .......... 

1. Grey shale .......... 

Section No. 2. Cayton Bay (Text-fig. 2), exposure in north face of 
Yons Nab headland. This section is complete, but like Section 
No. I is liable to be cut into by washouts infilled with deltaic 
sandstone (Moor Grit). Up to 10 ft. 7 ins. of sediment (G.L.S.) 
exposed. 

Upper Deltaic Series 
8. Moor Grit. 

Grey Limestone Series 

7. Chocolate-brown to grey coloured shale with fragmentary shells . i 5 

6. Fossihferous, chocolate-brown mudstone, variable in thickness. 

Glyptocythere scitula ; Vernoniella bajociana ; Vernoniella ? 
caytonensis ; Systenocythere ovata ; Paracytheridea ? caytonensis ; 
Progonocythere acuminata ; Progonocythere yonsnabensis and 
Praeschuleridea subtrigona intermedia ..... 2 o 

to 
9 

5. Grey fossihferous shale with : Glyptocythere scitula ; Vernoniella 

bajociana ; Fuhrbergiella [Praefuhrbergiella) horrida horrida ; 
Monoceratina scarboroughensis ; Praeschuleridea subtrigona inter- 
media ; Progonocythere yonsnabensis and Caytonidea faveolata . 2 o 

4. Brown fossiliferous mudstone with : Glyptocythere scitula and 

Vernoniella bajociana ........ o 4 

3. Grey shelly shale with : Glyptocythere scitula ; Vernoniella bajociana 

and Praeschuleridea subtrigona intermedia ..... 2 9 

2. Grey-black sulphurous shale ....... 2 i 

Middle Deltaic Series 
I. Flaggy yellow sandstone. 



82 R. H. BATE 

ft. in. 

Section No. 3. White Nab (Text-fig. 2), Scarborough. Exposure 

largely along the foreshore, cut out as a series of steps by the sea 

— ^well exposed and possibly almost complete, lowermost beds 

below Low Water Mark. 25 ft. 6 in. of marine sediment exposed. 

Upper Deltaic Series 
18. Massive deltaic sandstone (Moor Grit) forming base of high cliffs. 

Grey Limestone Series 

17. Sandy shale passing into base of Moor Grit above . . . i 

16. Fossiliferous, grey calcareous shale, partly ironstained. Fuhrber- 

giella (Praefuhrbergiella) horrida horrida . . . . o 7 

15. Fossiliferous shale with mudstone nodules. Internal casts, possibly 

of Glyptocythere scitula and Praeschnleridea subtrigonu intermedia 8 

14. Mudstone with Glyptocythere scitula ; V ernomella bajociana and 

Praeschideridea subtrigona intermedia ..... o 4 

13. Grey shale with ooliths ........ o 2 

12. Grey to chocolate-brown fossiliferous shale with Glyptocythere 
scitula ; V ernoniella bajociana and Praeschideridea subtrigona 
intermedia .......... i 

II. Ironstained mudstone crowded with the lamellibranch Gervillia 

scarburgensis ......... 5 

10. Sandy limestone containing large specimens of Gervillia scarburgen- 
sis Glyptocythere scitida and Praeschideridea subtrigona intermedia i 5 
9. Purplish-brown calcareous shale with Glyptocythere scitula ; V er- 
noniella bajociana and Praeschideridea subtrigona intermedia . i o 
8. Ironstained, sandy limestone. Vernoniella bajociana . . . 1 6 
7. Fossiliferous grey shale with Glyptocythere scitula ; Vernoniella 
bajociana ; Monoceratina scarboroughensis and Praeschuleridea 
subtrigona intermedia . ....... i 2 

6. Calcareous shale with mudstone bed at top which varies from 

o in. to 9 in. in thickness ....... i 5 

5. Massive, well bedded sandy limestone, fossihferous though no 
micro fauna so far obtained ....... 

4. Hard calcareous shale, almost an argillaceous limestone 
, 3. Soft, grey fossiliferous shale. Glyptocythere scitula ; Monoceratina 
scarboroughensis ; Fuhrbergiella (Praefuhrbergiella) horrida hor- 
rida and Praeschuleridea subtrigona intermedia .... 

2. Ironstained limestone ........ 

I. Grey sandy shale with large belemnites. Glyptocythere scitula ; 
Vernoniella bajociana and Praeschuleridea subtrigona intermedia 
seen to ......... . 



6 

I 


7 
6 


I 





I 






MIDDLE JURASSIC OSTRACODA 



83 



Section No. 4. Hundale point (Text -fig. 2) . A complete succession 
of these beds is exposed in the cliff face and along the foreshore, 
with a maximum thickness of 62 ft. 4 in. 

Upper Deltaic Series 
32. Moor Grit — carbonaceous and more thinly bedded at base. 



ft. 



in. 



Grey Limestone Series 
31. Grey, sandy and micaceous shale, ironstained 
30. Dark grey sulphurous shale ..... 
29. Fossiliferous mudstone, variable in thickness, maximum 
28. Grey shale, ironstained at base ..... 



27. Chocolate-brown, fossiliferous shale ...... 

26. Grey calcareous shale, almost an argillaceous Umestone. Glyptocy- 
there scitula ; Vernoniella bajociana ; Fuhrbergiella {Praefuhr- 
bergiella) horrida horrida ; Praeschuleridea suhtrigona intermedia . 

25. Chocolate-brown mudstone, shelly. Glyptocythere scitula ; Vernoni- 
ella bajociana and Monoceratina scarboroughensis 

24. Purplish brown, fossiliferous shale grading into bed 25. Glypto- 
cythere scitula ; Vernoniella bajociana ; Monoceratina scar- 
boroughensis and Praeschuleridea subtrigona intermedia 

23. Grey shale becoming less fissile towards base. Glyptocythere 
scitula ; Vernoniella bajociana ; Monoceratina scarboroughensis 
and Fuhrbergiella (Praefuhrbergiella) horrida horrida . 

22. Grey-black, very fossiliferous shale. Ostracod fauna at top of 
sequence : — Glyptocythere scitula ; Monoceratina scarboroughen- 
sis ; Fuhrbergiella [Praefuhrbergiella) horrida horrida and 
Praeschuleridea subtrigona intermedia. 

Fauna 7 ft. from base : — ostracods indet. 

Fauna 6 ft. from base : — Ljubimovella piriformis. 

Fauna 5 ft. from base : — Glyptocythere scitula and Vernoniella 
bajociana. 

Fauna 2 ft. from base : — Monoceratina scarboroughensis ; 
Cloughtonella rugosa and Praeschuleridea subtrigona intermedia. 

Fauna at base : — Glyptocythere scitula ; Ljubimovella pirifor- 
mis and Praeschuleridea subtrigona intermedia .... 

21. Calcareous shale. Cloughtonella rugosa ; Fuhrbergiella [Praefuhr- 
bergiella) horrida horrida ; Praeschuleridea subtrigona intermedia ; 
Monoceratina scarboroughensis ; Systenocythere ovata and Ljubi- 
movella piriformis . ........ 



3 




8 


I 




7 







4 





to 


7 







10 







10 







9 







4 



ir 



II 



R. H. BATE 



ft. in. 



20. Sandy limestone, very fossiliferous [Trigona sp. Pholadomya sp. 

etc.). Glyptocythere scitula ....... o 8 

19. Calcareous, grey shale with macrofossils as for bed 20. Glypto- 
cythere scitula ; Fuhrbergiella {Praefuhrbergiella) horrida horrida ; 
Praeschuleridea suhtrigona intermedia and Systenocythere ovata . 4 9 

18. Grey sandy shale. Systenocythere ovata ..... i 11 

17. Calcareous sandstone ........ i 5 

16. Dark grey shale extensively burrowed by marine organisms — the 

burrows infilled with sand. Belemnites present — no microfossils 2 10 

to 
3 o 

15. Hard calcareous shale. Pentacrinus ossicles common . . . i 2 

14. Grey sandstone with Pentacrinus ossicles. The Crinoid Grit. 
Current bedded with ripple markings along the bedding planes. 
Worm burrows common. This bed grades upwards into bed 15. 
Foraminifera but no ostracods have been obtained from this 
bed ........... 4 o 

to 
6 2 

13. Grey shale .......... 1 3 

12. Purplish-brown mudstone with Glyptocythere polita and Praeschuler- 
idea subtrigona intermedia ....... I 10 

II. Limestone Glyptocythere polita ; Malzia unicarinata and Prae- 
schuleridea subtrigona intermedia ...... I 3 

10. Sandstone .......... o 9 

g. Grey sandy shale. Glyptocythere polita ; Malzia bicarinata and 

Praeschuleridea subtrigona intermedia ..... 2 2 

8. Dark grey shale. Glyptocythere polita ; Vernoniella bajociana ; 
Fuhrbergiella {Praefuhrbergiella) horrida horrida and Prae- 
schuleridea s^ibtrigona intermedia ...... o 6 

7. Grey mudstone. Glyptocythere polita in this bed, as in bed 12, occurs 
in extremely large numbers. Other members of the ostracod 
fauna are : — Progonocythere acuminata ; Vernoniella bajociana 
QXid Praeschuleridea subtrigona intermedia. . . . . i i 

6. Grey shale. Glyptocythere polita and Praeschuleridea subtrigona 

intermedia .......... i 2 

5. Grey calcareous sandstone. Glyptocythere polita and Glyptocythere 

costata .......... 3 4 

4. Sandy shale ..........20 

3. Hard black shale ......... 2 7 

Middle Deltaic Series 
2. Massive sandstone ......... 11 7 



MIDDLE JURASSIC OSTRACODA 



85 



ft. 

seen to o 



I. Black shale ........ 

Section No. 5. Ravenscar (Text -fig. 2). As in the previous 
section, a complete sequence of marine strata is exposed, 
although the junction between the Grey Limestone Series and the 
Upper Deltaic Series is obscured by talus. The beds crop out 
high up in the steep cliff face, the Moor Grit of the Deltaic Series 
above capping the cliffs. 



m. 
2 



Upper Deltaic Series 

17. Soft yellow, false bedded sandstone, about 15 ft. of which cap 
the cliffs at this point. 

Grey Limestone Series 

16. Reddish-brown, rather soft sandstone above which there is 3-4 ft 
of sediment obscured by talus ..... 

15. Ochre coloured sandy shale ...... 

14. Ironstone with fossil casts ...... 

13. Ochre coloured sandy shale ...... 

12. Dark grey calcareous shale, fossiliferous throughout with shelly 
bands and nodules. 

Fauna 27 ft. from base : — Monoceratina scarboroughensis ; 
Ljuhimovella piriformis ; Fuhrbergiella {Praefuhrbergiella) horrida 
horrida and Praeschuleridea subtrigona intermedia. 

Fauna 25 ft. from base : — indeterminate ostracods. 

Fauna 21 ft. from base : — Glyptocythere scitula ; Vernoniella 
bajociana and Monoceratina scarboroughensis. 

Fauna 19 ft. from base : — indet. internal casts. 

Fauna 17 ft. from base : — indet. internal casts. 

Fauna 15 ft. from base : — Glyptocythere scitula ; Praeschuleridea. 
subtrigona intermedia and Fuhrbergiella {Praefuhrbergiella) horrida 
horrida. 

Fauna 13 ft. from base in shelly band : — Glyptocythere scitula. 

Fauna 11 ft. from base in shelly bed : — Glyptocythere scitula 
and Ljubimovella piriformis. 

Fauna 6 ft. from base : — Glyptocythere scitula ; Glyptocythere 
costata?; Ljubimovella piriformis ; Fuhrbergiella {Praefuhrbergiella) 
horrida horrida and Praeschuleridea subtrigona intermedia. 

In the shale sampled within the basal 4 ft. of shale, no microfauna 
has been so far obtained ....... 

II. Fossiliferous yellow sandstone with Pentacrinus ossicles. The 
Crinoid Grit . . . . 



I 


10 


• 3 


5 





4 


• 4 






30 



86 R. H. BATE 



ft. in. 



10. Light-grey, calcareous shale with abundant Gervillia scarburgensis . 
Fauna at top of bed : — Malzia unicarinaia ; Praeschuleridea 
suhtrigona intermedia ; Glyptocythere costata and Vernoniella 
bajociana. Fauna at base : — Malzia unicarinaia ; Malzia 
bicarinata ; Glyptocythere costata and Progonocythere acuminata ? 2 3 

9. Dark-grey, fossiliferous shale. Glyptocythere polita and Malzia 

unicarinaia .......... i 9 

8. Fossiliferous argillaceous limestone with Gervillia scarburgensis. 
Glyptocythere polita ; Glyptocythere costata ; Malzia unicarinaia ; 
Malzia bicarinata and Praeschuleridea subtrigona intermedia . o 4 

7. Grey sandy shale. Fauna at top : — Glyptocythere polita ; Malzia 
bicarinata ; Progonocythere acuminata and Praeschuleridea sub- 
trigona intermedia. Fauna at Base : — Glyptocythere polita ; 
Vernoniella bajociana and Praeschuleridea subtrigona intermedia . 2 6 

6. Calcareous mudstone, almost an argillaceous limestone. Glypto- 
cythere polita ; Progonocythere acuminata and Praeschuleridea 
subtrigona intermedia ........ i 2 

5. Rubbly bed — same lithology as bed 4. Glyptocythere polita and 

Praeschuleridea subtrigona intermedia ..... 7 

4. Dark grey, massive, sandy limestone. Glyptocythere polita and 

Praeschuleridea subtrigona intermedia ..... 2 

3. Dark-grey shale extensively burrowed by marine organisms — the 
whole bed being a mixture of shale and sandstone. Large speci- 
mens of Gervillia scarburgensis present. Glyptocythere polita and 
Praeschuleridea subtrigona intermedia ..... i 6 

2. Grey-black, rather brittle shale — no microfauna . . . . i 10 

Middle Deltaic Series 

I. Deltaic sandstone, massively bedded ..... 6 10 

Section No. 6. Hawsker (Text-fig. 2.). The thickness of the Grey 
Limestone Series has been considerably reduced and although the 
top of the succession is not exposed, the observed thickness of 
22 ft. 5 in. cannot be far short of the total in this area. 

Grey Limestone Series 

13. Well bedded grey sandstone, rather coarse grained, virtually a 
grit, which grades down into a chocolate-brown sandstone shelly 
in parts. Ostracods {Glyptocythere scitula ? and Praeschuleridea 
subtrigona intermedia) present though somewhat decalcified 

seen to 5 2 

12. Sandy limestone, crowded with shells towards the base — rather 
fissile on weathering. Vernoniella ? caytonensis ; Systeno- 
cythere ovata at base ........ 2 5 



MIDDLE JURASSIC OSTRACODA 87 

ft. in. 
II. Sandy and very fossiliferous chocolate-brown shale. Southcavea 
microcellulosa ; Praeschuleridea subtrigona intermedia ; Eocy- 
theropteron ? sp. ; Glyptocy there scittila ? and Fuhrbergiella 
{Praefuhrbergiella) horrida horrida ...... i 6 

10. Chocolate-brown shale. Glyptocytherescitula ; Vernoniellabajociana 

and Praeschuleridea subtrigona intermedia . . . . i 8 

9. Grey shale crowded with fossils at top. Glyptocythere scitula and 
Vernoniella bajociana at the top and Glyptocythere scitula ; 
Cloughtonella rugosa and Fuhrbergiella {Praefuhrbergiella) horrida 
horrida at base ......... 3 o 

■8. Grey shale with large nodules of limestone ..... 2 o 

7. Grey fossiliferous shale with Glyptocythere scitula ; Vernoniella 
bajociana ; Fiihrbergiella {Praefuhrbergiella) horrida horrida ; 
Ljubimovella piriformis ; Systenocythere ovata ; Progonocythere 
acuminata and Praeschuleridea subtrigona intermedia ■ • 3 

6. Calcareous sandstone, fossiliferous. No microfossils obtained. The 

Crinoid Grit of elsewhere ....... 2 o 

5. Dark-grey sandy shale, fossiUferous along bedding planes. Internal 

casts of Glyptocythere sp. . . . . . . .1 8 

Middle Deltaic Series 

4. Light-grey shale with sandstone lenses ..... 

3. Grey and white laminated sandy shale — sandstone lenses 
2. Grey sandy shale with i ft. i in. lens of sandstone at top. Shale 
carbonaceous with plant remains ...... 

I. Grey sandstone ....... seen to 

Section No. 7. Bloody Beck (Text-fig. 3). Of the inland exposures 
of the Grey Limestone Series, this is by far the best — it is complete and 
all beds are accessible along the course of the stream. 29 ft. 4 in. of 
marine sediment are developed here. 

Upper Deltaic Series 

25. Moor Grit — massively bedded sandstone forming the base of the 
Deltaic Series — flaggy towards base, grading into bed below. 

Grey Limestone Series 

24. Sandy, micaceous, dark-grey shale ...... 

23. Clay ironstone with specks of pyrite ...... 

22. Fossihferous grey, sandy shale, no microfossils .... 

21. Light-grey mudstone, very fossiliferous. Glyptocytherescitula 






8 


2 


I 


3 





4 


6 



2 


10 





7 


I 


2 





6 



R. H. BATE 



ft. in. 



20. Fossiliferous grey sandy shale — internal casts of Glyptocythere ? sp. 
and Vernoniella ? sp. Beds 24-20 have suffered extensive 
decalcification — fresh material if possible to obtain would 
probably yield a larger fauna ....... 4 

19. Hard, grey, calcareous shale. Ostracods fragmentary and in- 
determinate ......... I o 

18. Grey calcareous shale, fossiliferous. Glyptocythere scitula and 

Vernomella hajociana ........ 9 

17. Hard, grey, fossiliferous mudstone. Glyptocythere scitula and Ver- 
noniella bajociana. . . . . . . . . o 11 

16. Hard, grey, calcareous shale — quite fossiliferous although the 
ostracod fauna appears to be restricted to the single species, 
Vernoniella hajociana ........ i 7 

15. Grey, rather shelly shale, softer than bed 16. Ostracod fauna 

restricted to Fuhrbergiella {Praefuhrbergiella) horrida horrida . i 4 

14. Fossiliferous calcarous shale — poor ostrocod fauna. Glyptocythere 

scitula and Praeschuleridea subtrigona intermedia . . . o 9 

13. Hard, grey, calcareous shale /mudstone with sub-rounded quartz 
grains embedded particularly in upper part. Glyptocythere 
scitula .......... o 3 

12. Sparsely fossiliferous shale with mudstone nodules. No ostracods 

recovered .......... 10 

II. Grey fossiliferous shale. Fauna at top : — indet. 

Fauna 3 ft. 8 in. from top : — Glyptocythere scitula ; Systeno- 
cythere ovata and a fragment of Praeschuleridea subtrigona inter- 
media. 

Fauna 6 ft. 6 in. from top : — Glyptocythere scitula ; Vernoniella 
bajociana and Praeschuleridea subtrigona intermedia ... 8 o 

10. Grey, very fossiliferous shale. Cloughtonella rugosa and Praeschuler- 
idea subtrigona intermedia ....... o 7 

9. Calcareous shale grading upwards into bed 10. It is, however, a 
much more resistant bed to weathering — fossiliferous. Glypto- 
cythere scitula ; Progonocythere acuminata and ? Southcavea micro- 
cellulosa .......... o 7 

8. Hard, grey shale, sparsely fossiliferous. Fuhrbergiella {Prae- 
fuhrbergiella) horrida horrida and indet. ostracod fragments . i o 
7. Grey sandy shale, fossiliferous. Mesocytheridea howardianensis ; 

Glyptocythere scitula and Malzia bicarinata . . . . o 10 

6. Grey-black fossiliferous limestone becoming shaly towards base. 
Fauna at top in limestone : — Glyptocythere polita and Glypto- 
cythere scitula. Fauna in shaly beds at the base consists entirely 
of Glyptocythere polita which occurs in enormous numbers . .2 o 



MIDDLE JURASSIC OSTRACODA 89 

ft. in. 

5. Well jointed, grey limestone. Glyptocythere polita and Praeschuler- 

idea subtrigona intermedia . . . . . . . o 9 

4. Grey, fossiliferous shale. Glyptocythere polita ; Malziaunicarinata ; 

Progonocythere acuminata and Praeschuleridea suhtrigona inter- 
media. .......... I 4 

3. Grey, well jointed, limestone. Glyptocythere polita and Praeschuler- 

idea suhtrigona intermedia ....... o 8 

2. Grey sandy shale. Glyptocythere polita and Praeschuleridea suh- 

trigona intermedia ........ 1 3 

Middle Deltaic Series 

1. Massive sandstone, flaggy in places .... seen to 8 o 

Section No. 8. May Beck, stream section. Exposures poor and 
discontinuous, the ostracod fauna being almost entirely leached out. 
Section at NZ/8gooi5. 

Upper Deltaic Series 
10. Moor Grit — flaggy at base ..... seen to 12 

Grey Limestone Series 

9. Sandy limestone. Glyptocythere scitula and ? Fuhrhergiella {Prae- 

fuhrhergiella) horrida horrida ..... seen to 2 9 

8. Rubbly sandy bed with mudstone nodules ..... i 2 

7. Grey, calcareous sandstone. Praeschuleridea suhtrigona intermedia i i 

Section at NZ/890017, about 100 yds. downstream. 

6. Grey flaggy sandstone with fossil casts . . . seen to i o 
Section at NZ/891019, about 100 yds. downstream. 

5. Grey calcareous sandstone with fossil casts. This bed is probably 

more correctly a sandy limestone, leaching of the CaCOj reducing 

the bed to a sandstone. Praeschuleridea suhtrigona intermedia 3 3 

4. Purplish-red siltstone with decalcified shells. Glyptocythere scitula 

and Vernoniella bajociana ....... o 4 

3. Grey sandstone with fucoid markings . ..... o 7 

2. Grey, brittle shale^ — ^ostracod internal casts ..... i 7 
I . Light-grey shale with plant remains .... seen to 3 o 

Section No. 9. Ramsdale Beck, stream section. Only 9 ft. 6 in. of 
marine shale exposed in the left bank of the stream (facing downstream) . 
Although macrofossils are present within the shale, all the microfauna 
has been leached away except for the occasional indeterminate internal 
cast. 

GEOL. 11, 3 7 



go 



R. H. BATE 



Section No. io. Eller Beck, stream section, exposing 6 ft. 5 in. of 
marine sediment in the right bank of the stream, below road bridge. 

Grey Limestone Series 

5. Dark grey shale — sandy and ironstained. Internal casts of 
Glyptocythere ? sp. 

4. Calcareous mudstone with macrofossils. Glyptocythere scitula and 
Praeschuleridea subtrigona intermedia ..... 

3. Grey, sandy shale with a few fossils ...... 

2. Calcareous grey shale. This bed has a good calcareous cement and 
has not been so extensively decalcified. Glyptocythere scitula ; 
Vernoniella bajociana and Praeschuleridea subtrigona intermedia . 
Dark grey shale, internal casts of Glyptocythere ? sp., and Ver- 
noniella bajociana ...... seen to 



fi- 



tn. 



I. 



Section No. ii. Harland Beck, stream section in left bank in a 
disused bend of the stream just above the junction of the Harland Beck 
with the river Dove : 14 ft. 4 in. exposed. 

Grey Limestone Series 

7. Sandy shale. Glyptocythere scitula ; Vernoniella bajociana and 
Praeschuleridea subtrigona intermedia . . . seen to 

6. Grey, sandy limestone, fossiliferous along the bedding planes 
Praeschuleridea subtrigona intermedia .... 

5. Grey sandy shale with calcareous nodiiles .... 

4. Massive bedded sandstone almost a grit, flaggy at base . 

3. Sandy shale ......... 

2. Dark-grey shale, almost a mudstone in appearance. Lamellibranch 
Gervillia scarburgensis common. Glyptocythere scitula and Prae- 
schuleridea subtrigona intermedia ...... 

I. Dark-grey mudstone. Glyptocythere scitula and Praeschuleridea 
subtrigona intermedia ...... seen to 

Section No. 12. Bogmire Gill (Text-fig. 3), an almost complete 
stream section exposing some 40 ft. 3 in. of the Grey Limestone Series. 



Upper Deltaic Series 



10. Moor Grit — flaggy at base 



seen to 



Grey Limestone Series 



9. Grey sandy shale 






9 


3 


4 


10 


6 


2 


I 


I 






MIDDLE JURASSIC OSTRACODA 91 

ft. in. 

8. Dark-grey ironstained shale with fossils. Glyptocythere scitula and 

Vernoniella bajociana ........ i 6 

7. Grey-brown mudstone. Glyptocythere scitula ; Vernoniella bajo- 
ciana ; Progonocythere acuminata and Praeschuleridea subtrigona 
intermedia .......... 

6. Dark-grey, fossiliferous shale. Glyptocythere scitula and Vernoniella 
bajociana .......... 

unexposed section approx. 2 ft. 

5. Flaggy yellow sandstone with fossil casts ..... 

4. Grey sandy shale ......... 

3. Grey ironstained shale ........ 

unexposed section approx. 20 ft. 

2. Calcareous sandstone, known as the Crinoid Grit. Fossiliferous and 

false bedded. Ostracod internal casts, Glyptocythere ? sp. . . 10 

to 
12 

1. Steel-grey, flinty limestone, ripple marked along bedded planes. 

Indeterminate ostracods ..... seen to 3 

Section No. 13. (Text-fig. 3). A small, disused quarry on Yearsley 
Moor, exposing 8 ft. 3 in. of marine sediment. 

Upper Deltaic Series 

3. Yellow, rather soft, flaggy sandstone with plant remains along 

bedding planes ...... seen to approx. 6 

2. Yellow sandy bed with lenses of grey shale . . . . . i 

Grey Limestone Series 

I. Buff coloured sandy limestone, weathers to a sandstone — -fossili- 
ferous. 

Fauna approx. 3 ft. 6 in. from the base : — Glyptocythere scitula ; 
Praeschuleridea subtrigona intermedia ; Mesocytheridea howardian- 
ensis ; Fuhrbergiella [Praefuhrbergiella) horrida horrida and 
Systenocy there ovata. 8 

Section No. 14. (Text -fig. 3). Three isolated sections within the 
Grey Limestone Series, exposed in Stonecliff Wood, above the York- 
Scarborough railway-line. Altogether a maximum of 20 ft. i in. of 
sediment is exposed. 



92 R. H. BATE 



Grey Limestone Series 



ft. in. 



8. Sandy, fossiliferous limestone exposed at map reference SE/743676. 

Fauna at top of section : — Eocytheropteron ? sp. ; Southcavea 
microcellulosa and Praeschuleridea subtrigona intermedia. 

Fauna 3 ft. 8 in. from base : — Southcavea microcellulosa ; 
Praeschuleridea subtrigona intermedia ; Glyptocythere scitula ; 
Mesocytheridea howardianensis and Fuhrbergiella [Praefuhrber- 
giella) horrida horrida. Fauna i ft. 6 in. from base includes all 
those listed above, with the exception of Fuhrbergiella 
(Praefuhrbergiella) horrida horrida ...... 5 8 

Section at SE/744676 : — 

7. Grey limestone weathering to orange-red sandstone containing fossil 

casts. This bed is largely decalcified at outcrop . . . i 4 

to 
2 6 

6. Yellow sand, false bedded at top. Fossil casts . . . . i 7 

5. Limestone, impersistent laterally, being replaced by yellow sand. 
Glyptocythere scitula ; Eocytheropteron ? sp. ; Praeschuleridea 
subtrigona intermedia and Southcavea microcellulosa . . . i 8 

4. Soft yellow sandstone — easily weathers to a sand — fossil casts 

common in lower part ........ 3 o 

3. Shelly limestone, impersistent and replaced laterally by sand. 
Glyptocythere scitula and Southcavea microcellulusa 

2. Reddish -brown sand with fossil casts ...... 

1. Coarse silver sand ....... seen to 

Section at SE/740675 :— 

2. Grey limestone. Praeschuleridea subtrigona intermedia ; Meso- 

cytheridea howardianensis ; Glyptocythere scitula and Vernoniella 
bajociana. This bed is possibly at a slightly lower horizon than 
those listed in the two sections above ..... 9 

I. Yellow sandstone ....... seen to i 6 



III STRATIGRAPHY 

During Middle Jurassic times an important axis of downwarping, the Cleveland 
Axis, extended approximately east-west through the centre of the Yorkshire Basin, 
cutting the present coastline somewhere about Ravenscar (Text-fig. i). This axis 
of movement played an important role during the deposition of the Grey Limestone 
Series. 






8 





4 


2 


5 



MIDDLE JURASSIC OSTRACODA 93 

The Grey Limestone Series or Scarborough Beds are well exposed along the coast 
as far north as Whitby where the outcrop turns inland. In the northern part of the 
Yorkshire basin, the Cleveland Hills (Text-fig. 4), the outcrops are generally rep- 
resented by expanses of coarse calcareous grit known as the Crinoid Grit. Down the 
western part of the outcrop, exposures are sometimes good but incomplete ; the 
most complete inland exposures are found towards the centre of the basin in stream 
sections. 

Fox-Strangways (1892 : 236) was the first to subdivide this marine series on 
lithological terms. His divisions consisted of an upper shale division, a middle 
sandstone division and a lower limestone division. This arrangement was continued 
by Richardson (191 1). However, the subdivision of the Grey Limestone Series in 
this way, although basically correct, is a simplification of what really takes place. 
In fact failure to recognize facies changes resulted in Arkell (1933 : 221) making the 
following statement " Although the exact horizons of the ammonites recorded from 
the Scarborough or Grey Limestone are not known, it may be presumed that most if 
not all came from the lowest or limestone division ". The impure limestones 
developed in the Scarborough district belong to the upper shale division. The 
lower so called limestone division is not exposed at Scarborough and may not, in fact, 
extend as far south. As many of the ammonites recorded come from Scarborough, 
it is clear that they must come from the upper division. The examination of the 
ostracod faunas brought out this fact and at the same time made it possible to 
correlate the various sections more precisely than would have been possible on purely 
lithological evidence. 

The division of the Grey Limestone Series on lithological evidence is here aban- 
doned, instead two ostracod zones are recognized : a lower zone of Glyptocythere 
polita and an upper zone of Glyptocythere scitula, the type section for both being taken 
at Hundale Point — see Section 4. 

Glyptocythere scitula Zone 

The ostracod fauna associated with the index ostracod of this zone is more varied 
than that of the lower zone and possesses several species which are restricted in their 
geographical distribution. The typical faunal assemblage of this zone is as follows: — 

Glyptocythere scitula ; V ernoniella bajociana ; Monoceratina scarboroughensis ; 
Fuhrbergiella (Praefuhrbergiella) horrida horrida ; Ljubimovella piriformis ; 
Systenocy there ovata and Praeschideridea subtrigona intermedia. 

Of this fauna P. subtrigona intermedia ranges throughout the entire Grey Limestone 
strata and is not indicative of either zone. The two ostracods Vernoniella bajociana 
and Fuhrbergiella (P.) horrida horrida are similarly found to occur in sediments below 
this zone, but they are relatively uncommon at the lower horizon and are not con- 
sidered to be characteristic there. 

A number of ostracods, restricted to this zone, have a limited geographic distribu- 
tion, the palaeoecological considerations of which will be dealt with later. These 
are : — Cloughtonella rugosa ; Cytheropteron ? yonsnabensis ; Caytonidea faveolata ; 



94 R- H. BATE 

Mesocytheridea howardianensis ; Vernoniella ? caytonensis ; Paracytheridea ? 
caytonensis ; Progonocythere yonsnabensis and Southcavea microcellulosa. 

The ostracod Progonocythere acuminata is present in this zone, but is equally, if 
not more so, as common in the lower zone of G. polita and is of little value strati- 
graphically. 

Glyptocythere polita Zone 

The diagnostic fauna of this zone is : — G. polita, G. costata ; Malzia unicarinata 
and M. bicarinata. As mentioned above (see also Text-fig. 24) , several ostracods are 
present in both zones and need not be listed again. The above assemblage is charac- 
teristic of and restricted to this lower part of the Grey Limestone Series. 

In all cases the junction between the zone of G. polita and G. scitula is clear-cut. 
There is a very slight overlap of one or two species in some cases, but this is relatively 
insignificant, and where observed limited to a few feet of strata only. A few species, 
however, as mentioned above are not restricted to either zone, but range throughout 
the complete succession (Text-fig. 24). 

Along the coast the most southerly exposure of the Grey Limestone Series is in 
Gristhorpe Bay. Here the beds rapidly thicken to the observed maximum of 15 ft. 
6 in. at the north of the Bay in Yons Nab headland (Section i, Text-fig. 2). The 
marine sediments consist entirely of calcareous shales and mudstones both here and 
on the other side of the headland in Cayton Bay (Section 2, Text-fig. 2) where a 
maximum of 10 ft. 7 in. was observed. In this area the basal sandstone of the Upper 
Deltaic Series (here a rather incipient lateral equivalent of the massive Moor Grit 
further north) cuts down into the marine sediments as a number of well exposed 
washouts. These have been described and figured by Black (1928). 

At White Nab, Scarborough, the Series is again exposed (Section 3, Text-fig. 2) and 
has increased to an exposed maximum of 25 ft. 6 in. The shales are rather more 
strongly calcareous, and impure sandy limestones are developed. The ostracod 
fauna throughout the sequence, as in the previous two sections, is indicative of the 
G. scitula zone. It is possible that the lower zone of G. polita may be present under 
the sea. However this part of the section is never exposed even at low tide and 
must remain not proven for the present. 

North of Scarborough, at Hundale Point, Cloughton, the marine strata attain their 
maximum development of 62 ft. 4 in. (Section 4, Text-fig. 2). This is exactly the 
same thickness of sediment as measured for the Grey Limestone Series a few miles 
further north at Ravenscar (Section 5, Text-fig. 2). In both cases there is a very 
thick development of calcareous shale in the upper part, tending to be arenaceous at 
the top of Hundale, whilst at Ravenscar a bed of sandstone is developed. At 
Hundale there is still a remnant of the sandy limestone of the White Nab succession, 
present towards the lower part of the shale sequence. This limestone is not rep- 
resented at Ravenscar. At both localities the shale beds are followed by a thick bed 
of calcareous sandstone in which Pentacrimis ossicles are plentiful. This is the 
Crinoid Grit which is more typically developed further north. The ostracod fauna of 
the shale beds is indicative of the G. scitula Zone. No ostracods have been obtained 



MIDDLE JURASSIC OSTRACODA 95 

from the calcareous sandstone (Crinoid Grit) and it is taken to belong to the G. 
scitula Zone. As all the sediments below this bed at Hundale and Ravenscar belong 
to the G. polita Zone, the bed is here, a good marker horizon. 

The strata belonging to the G. polita Zone here reach their maximum development 
and consist of calcareous and sandy shales, some impure limestones and calcareous 
sandstones. Some mudstones are also present and these invariably are crowded with 
the ostracod Glyptocythere polita. 

Just south of Whitby the outcrop of the Grey Limestone Series turns inland away 
from the coast. However, an excellent exposure has been obtained in a stream sec- 
tion at Hawsker, or more precisely, Hawsker Bottoms (Section 6, Text-fig. 2). 
According to the section described by Fox-Strangways (1892 : 237) only 6 ft. 4 in. of 
sediment at the top of the section is unexposed at the present time. In this section 
the upper shale horizon, so well developed at Hundale and Ravenscar, is much thin- 
ner and a thick bed of fossiliferous sandstone and a bed of sandy limestone are 
introduced at the top. The sandstone is almost certainly laterally equivalent to bed 
16 present at the top of the shale horizon at Ravenscar, but here much more massive 
in character. The lateral equivalent of the Crinoid Grit (bed 6) is not very strongly 
developed here and beds possibly belonging to the G. polita Zone are very much 
reduced in thickness. The shale bed which probably represents this horizon (bed 5) 
has only produced extensively decalcified internal casts of Glyptocythere sp., so that 
conclusive evidence is at the moment lacking. Stratigraphically, however, there is 
good reason to consider bed 5 as representing the G. polita Zone with beds 6-13 
belonging to the G. scitula Zone. 

To the north and west of Whitby the marine sediments of the Series are poorly 
exposed and appear to consist almost entirely of coarse grained fossiliferous grits in 
which Pentacrinns ossicles are common. This lithofacies has been named the 
Crinoid Grit (Richardson 1911 : 195 and 197) and is laterally equivalent not only to 
the calcareous sandstone containing Pentacrimis ossicles at Hundale and Ravenscar 
but to practically the whole of the G. scitula Zone. However, shale and limestone 
horizons are still to be found in this northern area but are definitely subordinate to 
the arenaceous facies. 

Towards the centre of the depositional basin the Grey Limestone Series continues 
to exhibit a wide variety of lithofacies. At the Bloody Beck stream section (Section 
7, Text-fig. 3) calcareous shales and mudstones predominate with some impure lime- 
stones coming in towards the base in the zone of G. polita. The Crinoid Grit appears 
to be absent here and the junction of the two ostracod zones falls between beds 6 and 
7. Only a few miles to the north-west the majority of the sediments exposed in the 
May Beck section (Section 8) are quite arenaceous. In the west at Bogmire Gill 
(Section 12, Text-fig. 3) a large part of the section is again arenaceous and the Crinoid 
Grit (bed 2) is well developed and shows strong false bedding. The sediments in the 
Harland Beck section (Section 11) are also predominantly arenaceous. Apart from 
the Bloody Beck section where beds of G. polita age definitely occur, the only possible 
exposure of these beds is at Bogmire Gill where a flinty ripple marked limestone is 
probably to be correlated with this zone. So far no ostracods have been extracted 



96 



R. H. BATE 



from this lithology in a recognizable form. The basal limestone at Bogmire Gill was 
seen to be 3 feet, and is probably not very much thicker. The G. poUta Zone at 
Bloody Beck is 6 feet in thickness. Not only does this zone diminish considerably in 
thickness north and south away from the Hundale/Ravenscar area but it also thins 
westwards. The probable area covered by sediments of the G. polita Zone is shown in 
Text-fig 4. 

The most westerly outcrop of the Grey Limestone Series runs north /south through 
the Howardian Hills and consists of strongly arenaceous limestones interbedded with 
pure sand. In many cases decalcification of the limestone has resulted in the pro- 
duction of beds of sand. No complete section is exposed. Section 13 (Text-fig. 3) 
on Yearsley Moor exposes 8 ft. 3 in. of sandy limestone overlain by yellow sandstone 
of the Upper Deltaic Series. Further south at Stonecliff Wood, near Whitwell a 



NORTH 
SEA 



-Zone of G.poltta 
Zone of G.scitulo 




Fig. 4. Plan view of the marine basin during the deposition of the Grey Limestone Series, 
showing the probable maximum extent of the two faunal zones. 



MIDDLE JURASSIC OSTRACODA 97 

rather more complete section is exposed (Section 14, Text-fig. 3) . Here sandy lime- 
stones are seen to be not only interbedded with false-bedded sand but are also 
observed to pass laterally into red sandstone or sand. This strongly arenaceous 
facies of the Grey Limestone Series is also present in the east, south of Gristhorpe Bay. 
Here, however, not exposed at the surface. The Fordon No. i borehole proved 5 feet 
of sandy limestone with belemnites (Falcon & Kent i960 : 27). This facies is, 
therefore, continued around the western and southern perimeters of the Yorkshire 
basin whilst coarse grits are present along the northern edge. The shales and 
impure limestones are the major development towards the east and centre of the 
basin. 

IV CONCLUSIONS 

The Grey Limestone Series of north-east England is considered to represent the 
marine deposits of a broad embayment which cut into the deltaic sediments in Middle 
Jurassic times. 

The transgression of the sea at this point was the result of downwarping along the 
Cleveland Axis, the lowermost beds of the Series (zone of Glyptocythere poHta) being 
deposited only towards the centre of the basin (Text-fig. 4). The higher beds (zone 
of Glyptocythere scitula) extend over a wider area and in part rest directly upon deltaic 
sediments. The probable maximum extent of these higher beds is indicated in Text- 
fig- 4- 

That deposition of the marine sediments occurred in shallow water is evidenced by 
the presence of ripple markings, false bedding, worm burrows etc. The macrofauna 
is also generally indicative of shallow water conditions, which, together with a possible 
lowering of the salinity close to a delta may explain the almost complete absence of 
ammonites. The shore-line of the Series is indicated by the change in facies to a very 
sandy limestone and even to pure sand around the south and western boundaries of 
the outcrop. To the north, the whole series is more coarsely arenaceous, the detrital 
material being brought in by the delta. The present day perimeter of the Grey 
Limestone Series outcrop approximates closely to the original shore-line. 

As shown in Text -fig. 5, there are two prominent sandstone horizons which extend 
across the basin. Each reflects the influence of the northern delta. A third sand- 
stone, the deltaic Moor Grit brings the period of marine deposition to a close. This 
type of sedimentation with marine shales and limestones interfingered by marine 
deposited deltaic sandstones is suggestive of intermittently rising sea-level, see 
Dunbar & Rodgers (1958 : 85, text-fig. 35c). 

The ostracod fauna is typically shallow water benthos and in the majority of 
species present, appears to be largely independent of bottom facies. There is, how- 
ever, a typical shore-line fauna consisting of : — Southcavea microcellulosa ; Meso- 
cytheridea howardianensis ; Praeschuleridea sitbtrigona intermedia and Glyptocythere 
scitula. A number of other species such as Vernomella bajociana ; Systenocy there 
ovata and Fuhrbergiella (P.) horrida horrida also occur in this very sandy shore-line 
facies, although they are only poorly represented. Towards the centre of the basin in 
slightly deeper water, the characteristic fauna consists of (in the G. scitula Zone) : — 
Glyptocythere scitula ; Vernoniella bajociana ; Monoceratina scarboroughensts ; 



98 R. H. BATE 

Ljuhimovella piriformis ; Praeschuleridea subtrigona intermedia ; Cloughtonella 
mgosa ; Systenocythere ovata and Fnhrbergiella (P.) horrida horrida. A number of 
other species such as Caytonidea faveolata ; V ernoniella ? caytonensis ; Paracytheridea 
? caytonensis and Progonocythere yonsnabensis also occur but are rather restricted in 
their geographical range, reflecting an environment most probably restricted to the 
Cayton Bay region. The ostracod Progonocythere acuminata is only sporadically 
present at this horizon. In the G. polita Zone the fauna appears to have lived in a 



FORDON 1&2 



■S.E. 




~ NW. 



Fig. 5. Section across the Yorkshire Basin showing the facies changes of the Grey Lime- 
stone Series. Sections 1-6 are indicated together with information from the Fordon 
bore-hole. Vertical scale exaggerated. 

similar environment, with regard to bottom sediment and depth of water, as the 
above and consists of the following : — Glyptocythere polita ; G. costata ; Malzia 
bicarinata ; M. iinicarinata ; Progonocythere acuminata and Praeschuleridea sub- 
trigona intermedia. V ernoniella bajociana occurs in the upper part of this zone whilst 
Fuhrbergiella (P.) horrida horrida has only been recorded from a single locality. No 
definite shore-line fauna has been identified in this lower zone, due, no doubt to lack 
of exposures, although the ostracod Mesocytheridea howardianensis occurs in sandy 
shale at the base of the G. scitiila Zone in the Bloody Beck section, whilst a little 
higher up in the same section there is a single specimen possibly belonging to South- 
cavea microcellulosa. The presence of these ostracods here may indicate a shallowing 
of the water or a change in environment to their liking. It is not possible, however, to 
draw any further conclusions at this stage. 



V SYSTEMATIC DESCRIPTIONS 

Subclass OSTRACOD A Latreille 1806 

Order PODOCOPIDA Miiller 1894 

Suborder PODOCOPINA Sars 1866 

Superfamily CYPRIDACEA Baird 1845 

Family PARACYPRIDIDAE Sars, 1923 

Genus PARACYPRIS Sars 1866 

Paracypris bajociana Bate 



MIDDLE JURASSIC OSTRACODA 99 

For complete synonymy see Bate 1964 : 9. 

Remarks. Paracypris bajociana has been recorded from the Lincolnshire Lime- 
stone and from the Cave, Whitwell and Millepore Oolites of N.E. England (Bate 
1963, 1963(2 and 1964). Plumhoff (1963 : 18) records this species from beds of 
discites age and younger from Noith Germany. 

The only occurrence within the Grey Limestone Series is at the base of the Gris- 
thorpe Bay sequence where two specimens have been found. 

Superfamily GYTHERACEA Baird 1850 

Family BYTHOCYTHERIDAE Sars 1926 

Genus MONOCERATINA Roth 1928 

Monoceratina scarboroughensis sp. nov. 
(PI. I, figs. 1-12) 

Diagnosis. Monoceratina, with finely punctate, subquadrate to elongate cara- 
pace, slightly constricted just anterior of mid-dorsal region. 

HoLOTYPE. I0.1711, top of bed 22, Hundale Point, Cloughton. 

Paratypes. Io. 1712-23, from top and base of bed 22 and bed 25, Hundale 
Point ; bed 5, Cayton Bay and bed 12 (21 ft. and 27 ft. from base), Ravenscar. 

Description. Carapace subquadrate in outline, slightly constricted just anterior 
of mid-dorsal region, the more elongate dimorphs are considered to be the males. 
Greatest length of carapace through mid-point ; greatest height in anterior or 
posterior third ; greatest width in posterior third. The shallow constriction (sulcus) 
does not extend below mid-point, the ventro-lateral part of the carapace being 
noticeably swollen, particularly postero-ventrally. Dorsal margin straight with 
distinct, rounded cardinal angles. Ventral margin incurved medially ; anterior 
rounded ; posterior triangular with a short, straight or slightly concave postero- 
dorsal slope and a long, convex postero- ventral slope. Shell surface finely orna- 
mented with small, round puncta, arranged in longitudinal rows in the male dimorph. 
Valves almost equivalve : mid-ventrally the left valve slightly overlaps the right, 
whilst dorsally the right valve overlaps the left, the degree of overlap increasing 
towards the posterior cardinal angle. Muscle scars consist of a subvertical row of 
four rectangular adductor scars and two antero-dorsal scars situated below mid- 
length and below the shallow sulcus. Hinge in the left valve consists of the slightly 
downset mid-dorsal edge of the valve acting as a hinge-bar ; not seen in the right 
valve, but presumably consisting of a simple dorsal groove for the articulation of the 
left valve hinge. Duplicature not clearly seen, though there appears to be a narrow 
anterior vestibule developed in one paratype (I0.1716). 

Dimensions 

Holotype I0.1711, female carapace (PI. i, figs. 1-3), length o-6i mm. ; height 
0-32 mm. ; width 0-25 mm. 



loo R. H. BATE 

I0.1719, female right valve, length o-6o mm. ; height 0-35 mm. I0.1720, female 
left valve (PL i, fig. 12), length 0-55 mm. ; height 0-33 mm. I0.1721, male carapace 
(PI. I, figs. 9-11), length 0-70 mm. ; height 0-32 mm. ; width 0-27 mm. I0.1722, 
female carapace, length 0-52 mm. ; height 0-27 mm. ; width 0-22 mm. I0.1723, 
female carapace (PI. i, figs. 4-8), length 0-57 mm. ; height 0-31 mm. ; width 0-23 
mm. 

Remarks. Monoceratina scarboroughensis differs from all previously named 
species in outline (greatest length being through mid-point and not dorsal of mid- 
point as in the majority of cases), surface ornamentation of fine puncta coupled with 
the presence of dimorphism. 

Family PROGONOGYTHERIDAE Sylvester-Bradley 1948 

Subfamily PROGONOCYTHERINAE Sylvester-Bradley 1948 

Genus CAYTONIDEA nov. 

Diagnosis. Progonocytherinae, oval-rectangular in outline with well rounded 
anterior and posterior margins. Low eye swelling situated at anterior cardinal 
angle. Cardinal angles prominent, broadly rounded. Hinge antimerodont. Muscle 
scars consist of subvertical row of four adductor scars, rounded antero-dorsal 
antennal scar and rounded antero-ventral mandibular scar. Radial pore canals 
long, straight, few in number. Left valve larger than right. 

Type species. Caytonidea faveolata sp. nov. 

Remarks. Only a single species can be placed in the genus at the present time 
and this is typified by a strongly reticulate ornament. The possession of a well 
rounded oval-rectangular carapace with an antimerodont hinge, type A muscle scar 
arrangement and a distinct eye swelling identifies the genus Caytonidea from all other 
cytheracean genera. The genus (feminine) is named after the type locality, Cayton 
Bay. 

Caytonidea faveolata sp. nov. 
(PL I, figs. 13-14 ; PL 2, figs, i-io ; Text-figs. 6, 7) 
Diagnosis. Caytonidea, with strongly reticulate ornament of 5-6 sided pits. 
Holotype. Io. 1831, bed 5, Cayton Bay. 

Paratypes. Io. 1832-35, horizon and locality as above and from bed 4, Gris- 
thorpe Bay. 




Fig. 6. Hinge of right valve, Caytonidea faveolata sp. nov. Paratype, I0.1832, approx. X190. 



MIDDLE JURASSIC OSTRACODA loi 

Description. Carapace oval-subrectangular in outline with well rounded anterior 
and posterior margins, slightly concave mid-dorsal margin and antero-medially 
incurved ventral margin. The carapace is constricted slightly in the mid-dorsal 
region, and is noticeably swollen in the postero-dorsal region. Greatest length of 
carpace through mid-point, greatest height in the anterior third, greatest width in 
the posterior third. Shell surface ornamented by prominent 5-6 sided pits. Ventrally 
the reticulate ornament is somewhat subdued and is dominated by a series of longi- 
tudinal ridges. A low, smooth, eye swelling is situated at the anterior cardinal angle. 
Left valve larger than the right which it over-laps along the ventral margin and slightly 




V. •-.-. 



V^^^;:.iO:':->V-^v.';.::-;.-.^ 

Fig. 7. Muscle scars, Caytonidea faveolata sp. nov. Paratype, I0.1833, approx. X 250. 

in the region of the anterior cardinal angle. Hinge antimerodont, seen only in the 
right valve : 4 anterior and 5 posterior teeth observed ; median groove long and 
locellate. Duplicature of moderate width, the inner margin and line of concrescence 
coinciding. Radial pore canals long, straight and widelyspaced, yobservedanteriorly. 
Muscle scars with rounded antero-dorsal antennal scar (Type A). 

Dimensions 

Holotype I0.1831, carapace (PL 2, figs. 1-4), length 0-56 mm. ; height 0-33 mm. ; 
width 0-30 mm. 

I0.1834, carapace (PL i, figs. 13, 14 ; PL 2, figs. 5, 6), length 0-65 mm. ; height 
0-37 mm. ; width 0-33 mm. I0.1835, carapace length 0-56 mm. ; height 0-34 mm. ; 
width 0-27 mm. 

Remarks. Externally similar to Southcavea reticulata Bate (1964 : 27, pi. 10, 
figs. 3-14 ; pi. II, figs. 1-4), Caytonidea faveolata differs in the presence of an eye 
swelling, type A muscle scar arrangement as against type D, and in being less convex 
in dorsal view. The reticulate ornament of S. reticulata differs markedly from that of 
C. faveolata in the presence of strong punctation inside the shallow pits. 

Genus CLOUGHTONELLA nov. 

Diagnosis. Progonocytherinae with subquadrate carapace, virtually parallel- 
sided in dorsal view. Ventero-lateral border convex projecting downwards and 



102 R. H. BATE 

outwards in type species. Dorsal margin of larger left valve concave with upstanding 
cardinal angles. Posterior traingular ; anterior rounded with narrow marginal 
border. Hinge antimerodont. Duplicature of moderate width, inner margin and 
line of concrescence coinciding. Radial pore canals straight, few in number, widely 
spaced. Muscle scars not observed. 

Type species. Cloughtonella rugosa sp. nov. 

Remarks. Cloughtonella is very close to Aulacocythere Bate (1963 : 198) from 
which it probably evolved. The general morphological features of these two genera 
suggest a very close relationship. However, Cloughtonella can be distinguished by 
the absence of the horseshoe-shaped swelling of Aulacocythere and does not possess an 
eye swelling. 

At present only a single species can be definitely assigned to the genus : Cloughtonella 
rugosa sp. nov. However, the ostracod Procytheridea hoffmani Brand (1961 : 159, 
pi. I, figs. 1-8) is possibly congeneric although tending to be more oval in side view, 
with the dorsal margin of the left valve broadly convex, passing down to the extreme 
posterior without any break at the cardinal angle. The dorsal margin of the left 
valve, male dimorph, may be slightly concave, however. P. hoffmanni does not 
appear to belong to the genus Micropneumatocythere Bate (1963a : 29), to which many 
of the European procytherids belong, nor is it a true Procytheridea Peterson (1954 : 
171) which is a much more posteriorly acuminate genus. It is here tentatively 
considered to be congeneric with C. rugosa. The specimens of Procytheridea hoffmanni 
examined here were obtained from a sample of the romani Schichten, South Hannover, 
sent to me by Dr. F. Plumhoff, Erdol A. G., Wietze krs. Celle. The known range of 
P. hoffmanni is romani to blagdeni Zones, that of C. rugosa uncertain because of the 
almost comiplete absence of an ammonite fauna, but probably just below blagdeni 
Zone. 

The genus Cloughtonella (feminine) is named after the type locality Hundale Point, 
Cloughton Wyke. 

Cloughtonella rugosa sp. nov. 
(PI. 3, figs. 1-13 ; Text-figs. 8, 9) 

Diagnosis. Cloughtonella, with small subquadrate carapace. Dorsal margin 
medially concave in left valve. Ornamentation consists of prominent diagonal, 
rather irregular median ridge extending from postero-dorsal to antero-ventral 
region. Weak longitudinal ridges occur on either side of diagonal ridge. Ventro- 
lateral border of valves project slightly outwards and downwards, with longitudinal 
groove above, particularly prominent in male dimorphs. Hinge antimerodont. 
Muscle scars and radial pore canals not seen. 

HoLOTYPE. I0.2118, base bed 22, Hunsdale Point, Cloughton Wyke. 

Paratypes. Io. 2119-36, horizon and locality as above and from horizon 2 ft. 
from base of bed 22, Hundale Point ; from bed 10, Bloody Beck and from base of bed 
10, Hawsker. 



MIDDLE JURASSIC OSTRACODA 103 

Description. Carapace subquadrate in outline, more elongate in the male 
dimorphs. Greatest length of carapace extends through mid-point whilst the greatest 
height is in the anterior third. Greatest width in the posterior third, although there 
is only a slight increase in width posteriorly when compared with the width in the 
anterior part of the carapace. This is clearly seen in dorsal view, the carapace 
tending to be almost parallel-sided. Anterior broadly rounded, posterior triangular. 
Posterior marginal border narrow ; anterior border broad, directed obliquely back 
towards anterior cardinal angle. Dorsal margin concave in the left valve, convex in 
the right. Cardinal angles prominent in both valves. Ventral margin tends to be 
straight, the central part of the ventral surface being flattened, overhung on either 
side by the convex ventro-lateral margins, which in some specimens tend to project 
slightly outwards although not sufficiently developed to be termed alate. Left valve 
larger than the right which it overlaps along the ventral margin and strongly over- 
reaches along the dorsal margin. Terminally the left slightly overreaches the right 
with the possible exception of at the extreme posterior where there is no overreach or 




Fig. 8. Left side, female carapace, Cloughtonella rugosa sp. nov. Holotype, I0.2118, approx. 

X85. 

overlap. Shell surface ornamented by a series of low ridges, the dominant ridge 
running obliquely across the carapace from the postero-dorsal region to the antero- 
ventral region. This oblique ridge is rather irregular in outline and is bounded on 
either side by short, also irregular ridges which tend to be lateral below the main 
diagonal ridge and vertical above. Just above the ventro-lateral margin, which in 
some specimens also bears lateral ridges, there appears to be a shallow groove which 
gives the ventro-lateral extension of the carapace a pinched-up appearance particularly 
noticeable in the male dimorphs. The intensity of the ornamentation varies in each 
specimen but generally the impression given is of a wrinkled carapace. The ventral 
surface is ornamented by 3-4 longitudinal ridges on each valve. Hinge antimerodont. 
In the left valve the terminal sockets are separated by a short median ridge above 
which the shell slopes upwards to the dorsal margin. As a result there is virtually no 
accommodation groove developed. In the right valve only the anterior dentate 
element has been observed, bearing some 5-6 teeth. The median groove is poorly 
developed in the material examined but appears to be loculate. Inner margin and 
line of concrescence coincide, the duplicature being of moderate width. Radial pore 
canals almost imperceptible but can be made out as being short, straight and few in 
number. Muscle scars not observed. 



I04 



R. H. BATE 




Fig. 9. Left side, male carapace, Cloiightonella rugosa sp. nov. Paratype, I0.2119, approx. 

X95- 
Dimensions 

Holotype, I0.2118, female carapace (PL 3, figs. 1-4 ; Text-fig. 8), length 0-53 mm. ; 
height 0-34 mm. ; width 0-28 mm. 

I0.2119, male carapace (PL 3, figs. 5-7 ; Text-fig. g), length 0-55 mm. ; height 
0-28 mm. ; width 0-26 mm. I0.2120, male carapace (PL 3, figs. 12, 13), length 
o-6i mm. ; height 0-33 mm. ; width 0-26 mm. I0.2121, female left valve (PL 3, 
fig. 11), length 0-48 mm. ; height 0-32 mm. I0.2134, male carapace (PL 3, fig. 10), 
length 0-54 mm. ; height 0-29 mm. ; width 0-22 mm. I0.2135, female carapace 
(PL 3, figs. 8, 9), length 0-51 mm. ; height 0-32 mm. ; width 0-26 mm. I0.2136, 
male right valve, length o-6o mm. ; height 0-29 mm. 

Remarks. Cloughtonella rugosa is close to the genus Aulacocythere in outline but 
as mentioned under remarks for the genus does not possess the generic characters of 
the latter. The present species is also close to Procytheridea hoffmanni Brand, which 
can, however, be distinguished by the strongly arched dorsal margin in the left valve ; 
presence of a definite accommodation groove and in the more positive ornamentation. 
The two species are probably congeneric however. 

Genus FUHRBERGIELLA Brand & Malz 1962. 
Subgenus PRAEFUHRBERGIELLA Brand & Malz 1962 

Fuhrbergiella [Praefuhrbergiella) horvida Brand & Malz. 

Remarks. Brand & Malz (1962 : 19) described a new subgenus Praefuhrbergiella 
with Fuhrbergiella (P.) horrida as type species. Two subspecies were introduced : 
Fuhrbergiella [P.) horrida horrida having a range of romani to blagdeni Zones and 
Fuhrbergiella {P.) horrida bicostata typically developed in the Sonninien-Schichten 
[sowerbyi to grandis Zones) but also occurring in the Coronaten-Schichten (pinguis 
Zone) . The subspecies recorded here from the Grey Limestone Series is Fuhrbergiella 
(P.) horrida horrida. 

Fuhrbergiella {Praefuhrbergiella) horrida horrida Brand & Malz 

(PL 4, figs. 1-12) 

1962. Fuhrbergiella {Praefuhrbergiella) horrida horvida Brand & Malz : 19, pi. 4, figs. 33-37 ; 

pi. 5, fig. 46. 
1962. Fuhrbergiella [Praefuhrbergiella) horrida horrida Brand & Malz : Simon & Bartenstein : 

141, pi. 20, fig. 32 ; table 9. 



MIDDLE JURASSIC OSTRACODA 105 

Material. Seventy-two specimens examined from the Grey Limestone Series, 
of which the following have been registered in the British Museum collections : 
10.2109-17. 

Description. Carapace subquadrate narrowing towards the posterior. Sexual 
dimorphism indicated by the presence of more elongate dimorphs, considered to be 
the males. Greatest length through mid-point ; greatest height in the anterior 
third ; greatest width in the posterior third. Shell surface strongly reticulate , 
the reticulae in adults being extended into thin lamellae. Postero-dorsally a keel- 
like extension of the carapace projects above the dorsal margin of the valve bending 
down at about mid-point to die out close to the anterior margin below mid-length. 
Along the ventral surface a ridge is developed which turns upwards anteriorly 
towards an anterior vertical ridge which bounds the broad, flattened anterior border, 
to die out below the prominent eye node. The latter is situated just below the 
prominent, well rounded anterior cardinal angle. A short ventro-lateral ridge may 
be present above the ventral ridge in some specimens. Posterior marginal border 
also well developed. Anterior and posterior margins may possess shor't spines. Left 
valve larger than the right which it overlaps along the ventral margin and over- 
reaches along the antero-dorsal and postero-dorsal slopes. Hinge antimerodont with 
strongly loculate terminal sockets in the left valve and a long denticulate median 
bar. Accommodation groove poorly developed. In the right valve the hinge 
consists of some 8 anterior teeth and approximately 7 posterior teeth. Median 
groove poorly preserved in the present material. Inner margin and line of con- 
crescence coincide, the duplicature both anteriorly and posteriorly being quite broad. 
Radial pore canals long, straight and widely spaced ; 8-9 anteriorly and 4 posteriorly. 
The selvage forms a prominent ridge around the free margin, outside of which there is 
a naxro'w flange developed around the anterior margin and along the ventral margin. 
Only the 4 oval adductor scars have been seen in the present material, the anterior 
muscle scars not being preserved. 

Dimensions 

I0.2109, female left valve (PI. 4, figs. 1-3), length 078 mm. ; height 0-45 mm. 
I0.2110, female carapace (PL 4, figs. 11, 12), length o-68 mm. ; height 0-37 mm. ; 
width 0-38 mm. I0.2111, male carapace (PI. 4, figs. 6-9), length 074 mm. ; height 
0-37 mm. ; width 0-37 mm. I0.2115, female left valve, length o-6i mm. ; height 
0-35 mm. I0.2116, female right valve (PI. 4, figs. 4, 5), length 075 mm. ; height 
0-40 mm. I0.2217, female right valve (PL 4, fig. 10), length 073 mm. ; height 0-38 
mm. 

Remarks. All the specimens of Fuhrhergiella present within the Grey Limestone 
Series are here referred to Fuhrhergiella (Praefuhrbergiella) horrida horrida although 
in some cases the presence of a short ventro-lateral ridge indicates some aflinity with 
the subspecies F. [P.) horrida bicostata Brand & Malz (1962 : 21, pi. 4, figs. 38-40). 
The variation observed is here, however, restricted to the subspecies F. [P.] horrida 
horrida. 

GEOL. II, 3 g 



io6 R. H. BATE 

Genus GLYPTOCYTHERE Brand & Malz 1962 

Glyptocythere costata sp. nov. 

(PL 5, figs. 1-7) 

Diagnosis. Glyptocythere with subquadrate, posteriorly tapered carapace. 
Ornamented medially by branching transverse ridges radiating outwards from dorsal 
margin ; ventro-laterally by single longitudinal ridge and ventrally by second 
longitudinal ridge. 

HoLOTYPE. I0.1775, base bed 10, Ravenscar. 

Paratypes. Io. 1776-82, from top and base of bed 10, and from bed 8, Ravenscar. 

Description. Carapace subquadrate, tapering towards the narrowly triangular 
posterior. Elongate dimorphs indicate the presence of males within the population. 
The shell surface is strongly ornamented by irregular, branching transverse ridges 
which radiate downwards from the dorsal margin of the valve and by two prominent 
longitudinal ridges. The uppermost ridge extends along the ventro-lateral part of 
the carapace whilst the lower ridge, being an increased development of one of the 
finer longitudinal ridges which occur on the ventro-lateral and ventral surfaces, is 
ventral in position and in some specimens forms a prominent ventral keel. Anterior 
and posterior with narrow marginal border. Left valve larger than the right which it 
overlaps along the ventral margin and overreaches around the anterior margin, in the 
region of the anterior cardinal angle and along the postero-dorsal slope. Greatest 
length through mid-point ; greatest height in the anterior third ; greatest width in 
the posterior third. Anterior broadly rounded, posterior narrow, triangular, with 
short, concave, postero-dorsal slope and convex postero-ventral slope. Ventral 
margin incurved medially. Dorsal margin strongly convex in the right valve con- 
cave in the left. A low eye swelling is suggested in the right valve only, elongate in 
outline and situated below the anterior cardinal angle. Internal characters not 
observed. 

Dimensions 

Holotype, I0.1775, female carapace (PI. 5, figs. 1-4), length 0-65 mm. ; height 
0-40 mm. ; width 0-37 mm. 

I0.1776, female right valve (PI. 5, fig. 7), length 0-62 mm. ; height 0-35 mm. 
I0.1777, female carapace, length o-6o mm. ; height 0-38 mm. ; width 0-36 mm. 
Io. 1780, male carapace, length o-8o mm. ; height 0-43 mm. ; width 0-42 mm. I0.1782, 
male carapace (PI. 5, figs. 5, 6), length (broken) 0-59 mm. ; height 0-34 mm. ; width 
0-32 mm. 

Remarks. So far this species has only been found in sediments of the Grey 
Limestone Series as exposed at Ravenscar. Glyptocythere costata resembles Glypto- 
cythere dorsicostata Brand & Malz (1962a : 145, pi. 21, fig. 10, table 9) although it 
differs in being smaller with a more pronounced posterior taper and in the absence of 
the dorso-median ridge which characterizes the latter. G. costata is also close to 



MIDDLE JURASSIC OSTRACODA 107 

Progonocythere jiiglandica (Jones, 1884), Sylvester-Bradley (1948 : 193, pi. 12, figs. 
5, 6, pi. 13, fig. 8 ; text-fig. 4) to which it could be ancestral. G. costata, however, 
differs in being smaller and in the possession of strong longitudinal ridges in the ventral 
and ventro-lateral regions. [It is here considered that the species Progonocythere 
jiiglandica should be assigned to the genus Glyptocythere, to which it bears greater 
relationship than to Progonocythere. This will, however, be discussed in more detail 
in a subsequent paper.] 

Glyptocythere polita sp. nov. 
(PL 5, figs. 8-1 1 ; PL 6, figs. 1-9) 

Diagnosis. Glyptocythere with subquadrate/subtriangular carapace. Shell sur- 
face smooth with occasional wrinkles in dorso-median part. Downwardly projected 
ventro-lateral margin may be extended into thin keel, particularly in female dimorph. 

HoLOTYPE. I0.1724, bed 7, Hundale Point, Cloughton. 

Paratypes. Io. 1725-49 and I0.2210, beds, 7 and 8, Hundale Point, Cloughton ; 
beds 6, 7 and 9 Ravenscar and bed 6 Bloody Beck. 

Description. Carapace subquadrate to subtriangular in outline with strong 
sexual dimorphism : the males being much more elongate in outline and quite strongly 
convex in dorsal view. Greatest length of carapace through mid-point with the 
greatest height in the anterior third. Greatest width just behind valve middle. 
Left valve larger than the right which it overlaps along the ventral margin and 
overreaches in the region of the antero-, and postero-dorsal slopes and slightly around 
the anterior margin. Shell surface smooth with a shallow median sulcus marking the 
position of the adductor muscle scars. Occasionally there is apparent a slight 
wrinkling of the dorso-median part of the carapace, but this is rarely well developed. 
Normal pore canals are large, circular and prominently displayed over the valve 
surface. The ventral surface of each valve is strongly ornamented with longitudinal 
ridges some of which may be bifurcate, about 4-6 per valve, the outermost being 
situated along the convex ventro-lateral extension of the carapace and may be 
developed as a thin, blade-like keel. One or two weaker longitudinal ridges may 
occur outside this keel on the ventro-lateral part of the carapace. The keel, when 
developed appears to be restricted to the female dimorphs, being little more than a 
well developed ridge in the males. Anterior margin of carapace broadly rounded with 
flattened marginal border. Posterior triangular with marginal border, concave postero- 
dorsal slope, almost straight in the male, and convex postero-ventral slope. Ventral 
margin medially incurved, sweeping upwards posteriorly in female dimorph. Ventro- 
lateral margin extended below ventral surface, also sweeping strongly upwards 
posteriorly in the female dimorph. Dorsal margin medially concave in the left valve 
with prominent cardinal angles ; in the right valve medially convex, although the 
strong median convexity noticeable here is really dorso-median in position. Hinge 
entomodont : left valve with terminal loculate sockets and a strongly dentate median 
bar, especially so antero-medially. Accommodation groove virtually absent. Right 



io8 R. H. BATE 

valve hinge complementary to the left. Hinge in juveniles antimerodont. Muscle 
scars consist of a subvertical row of 4 oval adductor scars with (as seen in a single 
specimen) a crescent-shaped antero-dorsal antennal scar. Mandibular scar not 
observed. This muscle scar type is placed in type A (Bate 1963 : 181) rather than 
type B. Inner margin and line of concrescence coincide the duplicature being of 
moderate width. Radial pore canals straight and widely spaced, approximately 9 
anteriorly and 4 posteriorly. Anteriorly a narrow flange may be developed outside 
the selvage but is rarely preserved. 

Dimensions 

Holotype, I0.1724, female carapace (PI. 6, figs. 1-4), length 0-84 mm. height 0-52 
mm. ; width 0-47 mm. 

I0.1725, male carapace (PI. 6, figs. 5-8), length i-i8 mm. ; height 0-53 mm. ; 
width 0-55 mm. I0.1736, female carapace (PL 5, fig. 10), length 0-71 mm. ; height 
0-46 mm. ; width 43 mm. I0.1737, female carapace (PL 6, fig. 9), length 0-85 mm. ; 
height 0-54 mm. ; width 0-45 mm. I0.1738, female left valve (PL 5, fig. 9), length 
0-82 mm. ; height 0-49 mm. I0.1739, female right valve, length 077 mm. ; height 
0-46 mm. I0.1741, female left valve (PL 5, fig. 11), length 0-72 mm. ; height 0-46 
mm. I0.1742, male carapace, length i-oo mm. ; height 0-50 mm. ; width 0-51 mm. 
I0.1743, female carapace (PL 5, fig. 8), length 0-93 mm. ; height 0-56 mm. ; width 
0-48 mm. I0.1744, female carapace, length 0-71 mm. ; height 0-45 mm. ; width 
0-37 mm. 

Remarks. In general appearance Glyptocythere polita is similar to Progonocythere 
acuminata sp. no v., although it is not so strongly tapered posteriorly. The main 
differences between these two species are to be found in the dorsal margin, that of 
P. acuminata being straight and not convex as in G. polita. The dorsal margin of the 
left valve is also almost straight, or may be slightly concave, but not strongly so as in 
the present species. The much straighter dorsal margin of species of Progonocythere 
and the strongly convex dorsal development of the right valve in species of Glypto- 
cythere serve to distinguish the species of these respective genera. 

Glyptocythere scitula sp. nov. 
(PL 7, figs. 1-13 ; PL 8, figs. 1-9 ; PL 9, figs. 1-4 ; Text-fig. 10) 

Diagnosis. Squat, subquadrate to elongate Glyptocythere with strong branching 
and anastomosing ridges, radiating from dorsal margin. Ventro-lateral and ventral 
surfaces ornamented with longitudinal ridges, some of which bend upwards antero- 
ventrally. All ridges with rounded surfaces. Internal characters as for genus. 

Holotype. I0.1750, bed 5, Cayton Bay section. 

Paratypes. Io. 1751-74, from bed 5 Cayton Bay ; bed 5, Gristhorpe Bay ; beds 
17 and 18, Bloody Beck ; 6 ft. from base and 15 ft. from base bed 12, Ravenscar and 
from bed 10, Hawsker. 

Description. Carapace subquadrate in the female dimorph, elongate in the male. 



MIDDLE JURASSIC OSTRACODA 109 

Greatest length through mid-point ; greatest height in the anterior third. Greatest 
width in the posterior third. Carapace strongly convex in dorsal view, slightly 
constricted medially. Anterior and posterior with flattened marginal borders. Left 
valve larger than the right which it overlaps most strongly mid-ventrally and over- 
reaches along the anterior, posterior and dorsal margins. The degree of overreach 
is most strongly developed mid-dorsally. Anterior margin broadly rounded ; pos- 
terior triangular with concave postero-dorsal slope and convex postero-ventral slope. 
Ventral margin medially incurved ; dorsal margin strongly concave in the left valve 
convex in the right. The dorsal margin in the left valve besides projecting above that 
of the right valve is also very much thicker, almost keel-like. Ornamentation of 
carapace strongly developed and consists of transverse ridges which radiate outwards 
and downward from the dorsal margin, branching and anastomosing to produce a 
coarse reticulate pattern. The ridges are rounded and not lamellate. Ventro- 
lateral part of carapace strongly convex and ornamented by approximately 4 
longitudinal ridges. The same number of longitudinal ridges also occur along the 
ventral surface of each valve. The ventro-lateral ridges tend to turn upwards 
antero-ventrally. In some specimens the ornament is only poorly developed, here 
the very large, circular normal pore canals are particularly evident, widely scattered 
over the shell surface. Hinge entomodont : left valve with terminal loculate sockets 
and a strongly dentate median bar of which the anterior part is more coarsely dentate. 
Accommodation groove virtually absent, the valve sloping upwards to the dorsal 
margin from the base of the median bar. In the right valve 6 anterior and 6 posterior 
teeth are developed. Median groove expanded anteriorly, strongly loculate. Muscle 
scars (Type A) consist of a subvertical row of 4 oval adductor scars, a small round, 








■fll^ ^^■■^,^;U;' 




■:.^^§B^'M^M 






Fig. 10. Muscle scars, Glyptocythere scitula sp. nov. Female paratype, I0.1770, approx. X300. 

antero-dorsal antennal scar and a much larger rounded antero-ventral mandibular 
scar which appears as a rosette of several smaller scars. Inner margin and line of 
concrescence coincide the dupUcature being of moderate width. Radial pore canals 
long, straight, 9 anteriorly and 4 posteriorly. A n-A.rrow flange may be present around 
the anterior margin outside the selvage, whilst in the right valve a short ventral 
" lip " occurs just below the ventral incurvature of the valve. 



no R. H. BATE 

Dimensions 

Holotype, I0.1750, female carapace (PL 8, figs. 1-4), length 072 mm. ; height 
0*47 mm. ; width 0-43 mm. 

lo. 1751, male left valve (PL 7, figs. 11, 12), length 0-82 mm. ; height 0-45 mm. 
lo. 1752, female carapace (PL 8, figs. 6-9), length 0-77 mm. ; height 0-50 mm. ; 
width 0-47 mm. I0.1753, male carapace (PL 9, figs. 1-4), length 0-93 mm. ; height 
0-54 mm. ; width 0-52 mm. I0.1754, female left valve (PL 8, fig. 5), length 075 mm. ; 
height 0-48 mm. I0.1756, female right valve, length 0-63 mm. ; height 0-36 mm. 
lo. 1760, female right valve (PL 7, figs, i, 7), length (broken) 072 mm. ; height 
0-40 mm. I0.1768, female right valve (PL 7, figs. 4, 6, 10), length o-6o mm. ; height 
0*35 mm. I0.1769, female left valve (PL 7, figs. 5, 9), length o-6i mm. ; height 0-37 
mm. I0.1771, female right valve (PL 7, figs. 3, 8), length 0-68 mm. ; height 0-37 mm. 
ro.1772, male right valve (PL 7, fig. 13), length 0-82 mm. ; height 0-40 mm. I0.1773, 
female left valve, length 0-82 mm. ; height 0-48 mm. 

Remarks. Glyptocythere scitula although smaller than Glyptocythere tuberodentina 
Brand & Malz (1962a : 143, pi. 21, figs. 11, 12 ; table 9) is similar in general appear- 
ance. The two species may, however, be distinguished by the ornamentation which 
in G. tuberodentina consists essentially of a reticulate ornament on the lateral surface 
with prominent longitudinal ridges ventrolaterally. Both the longitudinal ridges and 
the ridges which comprise the reticulations are thin and lamellate. In G. scitula the 
ornament as described consists of radiating transverse ridges which produce a coarse 
reticulation on branching and anastomosing. In this species the longitudinal ridges 
and the transverse ridges are rounded, contrasting strongly with the lamellate ridges 
of G. tuberodentina. 

The known range of G. tuberodentina is from the middle of the acris Zone to the top 
of the friederici-augusti Zone of the Parkinsoni-Schichten. The range of G. scitula 
is more difficult to give precisely but probably does not occur higher than the blagdeni 
Zone of the Coronaten-Schichten. 

Genus MALZIA nov. 

Named after Dr. Heinz Malz of the Senckenberg Museum, Frankfurt-am-Main, 
Germany. 

Diagnosis. Progonocytherinae with subquadrate carapace, tapering to posterior. 
Anterior and posterior with flattened marginal borders. Ventro-lateral part of 
carapace extended into one or two keel-like projections. Low eye swelling developed 
at anterior cardinal angle. Speciesmay be dimorphic. Hinge entomodont. Radial 
pore canals long and straight, approximately 8 anteriorly, 3 posteriorly. Muscle 
scars as for subfamily (Type A) . 

Type Species. Malzia bicarinata sp. nov. 

Remarks. The genus Malzia (feminine) is erected here with two species : M. 
bicarinata having two ventro-lateral keel-like extensions and M. unicarinata sp. nov., 
having but a single valvular extension. It is considered that Malzia has a position 



MIDDLEJURASSICOSTRACOD\ in 

transitory between Progonocythere Sylvester-Bradley (1948 : 189) and Marslatourella 
Malz (1959: 19). This is suggested by the general similarity of carapace outline, 
muscle scars and radial pore canals present in all three. Progonocythere has a strong 
entomodont hinge and only a faint suggestion of an eye swelling. In Marslatourella 
the hinge is antimerodont and a strong eye tubercle is developed. At the same time 
prominent ventro-lateral outgrowths of the carapace occur. Malzia possesses a 
hinge which can be considered as entomodont, a low eye swelling and ventro-lateral 
outgrowths of the carapace. It appears, therefore, that the development of ventro- 
lateral outgrowths coupled with a change from an entomodont to an antimerodont 
hinge and the development of an eye tubercle results in the appearance in the 
Bathonian of the genus Marslatoiirella. This evolutionary series commences in the 
Bajocian with the genus Progonocythere an offshoot of which produces Malzia. This 
second lineage then continues giving rise to Marslatourella. 

The genus Marslatourella described by Malz from the Bathonian of France (Mars-la 
Tour and Boulonnais) and Germany (Eichberg) is also common in the Bathonian 
sediments of England. Species of this genus will be described in forthcoming pub- 
lications. 



Malzia bicarinata sp. nov. 

(PI. 9, figs. 5-8 ; PL 10, figs. 1-3 ; Text-figs. 11-14) 

Diagnosis. Malzia with two, short, well developed ventro-lateral keels. Internal 
details as for genus. 

HoLOTYPE. I0.1797, bed 9, Ravenscar section. 

Paratypes. I0.1798-1800, beds 7 and 8, Ravenscar and bed 7, Bloody Beck. 






J 




12 
Fig. II. Muscle scars. Malzia bicarinata sp. nov. Paratype, I0.1800, approx. X290. 

Figs. 12-13. Dorsal and left views, complete carapace Malzia bicarinata sp. nov. Holotype, 

lo. 1 797, approx. X85. 



112 R. H, BATE 

Description. Carapace subquadrate in outline with straight or very shghtly 
convex dorsal margin. Cardinal angles well developed. Anterior rounded ; 
posterior triangular with concave postero-dorsal slope and convex postero-ventral 
slope. Ventral margin medially incurved. Anterior and posterior with flattened 
marginal borders. Shell surface laterally smooth with two, stubby keels developed, 
the uppermost of which tends to project slightly outwards, particularly noticeable in 
dorsal view. Ventral surface may be smooth or possess two longitudinal ridges on 
each valve. A low eye swelling is situated just below the anterior cardinal angle, 
particularly noticeable in the nght valve. Greatest length of carapace through mid- 
point ; greatest height in the anterior third ; greatest width in the posterior third. 
Left valve larger than the right which it overlaps mid-ventrally and slightly over- 
reaches around the posterior and along the dorsal margin. Hinge entomodont, only 
seen in the left valve where the terminal loculate sockets are separated by a dentate 
median bar the dentition of which increases in coarseness towards the anterior. 
Accommodation groove narrow, elongace. Muscle scars (Type A), the antennal scar 
being round and antero-dorsal in position. Mandibular scar not seen. Radial pore 
canals straight and simple : approximately 8 anteriorly and 3 posteriorly. Duplica- 
ture in completely seen in present material. Selvage prominent external to which 
there is a narrow flange developed around the anterior margin and along the ventral 
margin. 




Fig. 14. Dorsal view, left valve hinge, Malzia bicarinata sp. nov. Paratype, I0.1799, approx. 

X105. 

Dimensions 



Holotype, I0.1797, carapace (PL 9, figs. 5-8 ; Text-figs. 12-13), length 070 mm. ; 
height 0-43 mm. ; width 0-41 mm. 



MIDDLE JURASSIC OSTRACODA 113 

I0.1798, carapace (PI. 10, fig. i), length 0-69 mm. ; height 0-43 mm. ; width 0-38 
mm. I0.1799, left valve (PI. 10, figs. 2, 3 ; Text-fig. 14), length 070 mm. ; height 
0-43 mm. 

Remarks. Malzia bicarinata cannot easily be confused with Marslatourella 
exposita Malz (1959 : 20, text-figs. 1-4), lacking the prominent eye tubercle and 
antimerodont type hinge of the latter. It is, however, sufficiently close as to be 
considered ancestral to M. exposita. The present species differs from species of 
Progonocythere in the development of ventro-lateral O'^itgrowths of the carapace and 
in the development of an eye swelling. Although somt species of Progonocythere may 
show the development of an eye swelling it is not so prominent as in Malzia although 
even here it is hardly a dominant feature of the carapace. 

Malzia unicarinata sp. nov. 
(PL 10, figs. 4-10 ; PI. II, lig. 1-4 ; Text-fig. 15) 

Diagnosis. Malzia, with single ventro-lateral keel. Dimorphic. 

HoLOTYPE. I0.1801, bed 9, Ravenscar section. 

Paratypes. Io. 1802-6, beds 8 and lo, Ravenscar and bed 4, Bloody Beck. 




Fig. 15. Right side, female carapace, Malzia unicarinata sp. nov. Holotype, I0.1801, approx. 

X105. 

Description. Carapace subquadrate in outline in the female dimorphs, elongate 
in the male. Ventro-lateral part of carapace convex, extended as a short, rather 
stubby keel. Greatest length through mid-point ; greatest height in the anterior 
third ; greatest width in the posterior third. Shell surface punctate. Anterior 
broadly rounded ; posterior broadly triangular with concave postero-dorsal slope and 
convex postero-ventral slope. Ventral margin medially incurved. Dorsal margin 
very slightly convex in the female, straight in the male although slightly concave 
just behind the anterior cardinal angle. Cardinal angles prominent, the anterior 
angle being broadly rounded with the posterior angle tending to be more acute. 
Anterior and posterior margins with well defined borders. Eye swelling elongate in 



114 R- H. BATE 

outline situated below the anterior cardinal angle, more prominently developed in 
the male dimorph. Left valve larger than the right which it overlaps mid-ventrally 
and overreaches along the remaining part of the ventral margin and along the dorsal 
margin. Internal details not seen. Radial pore canals as seen from the exterior 
straight and widely spaced, approximately 8 anteriorly and 3 posteriorly. A narrow 
flange extends around the anterior margin and although not clearly seen probably also 
along the ventral margin. 

Dimensions 

Holotype I0.1801, female carapace (PI. 10, figs. 4-8 ; Text-fig. i5),lengtho-7imm. ; 
height 0-42 mm. ; width 0-41 mm. 

I0.1802, female carapace (PL 10, figs. 9, 10), length 0-73 mm. ; height 0-44 mm. ; 
width 0-43 mm. I0.1804, female carapace, length 0-75 mm. ; height 0-46 mm. ; 
width 0-43 mm. I0.1806, male carapace (PL 11, figs. 1-4), length 0-85 mm. ; height 
0-45 mm. ; width 0-44 mm. 

Remarks. Malzia unicarinata is distinguished easily from M. bicarinata by the 
possession of but a single ventro-lateral keel. The male dimorph of M. unicarinata 
is, however, similar to Progonocythere acuminata sp. nov., from which it can be 
identified by the mid-laterally swollen, strongly convex carapace, a feature which 
characterizes species of this genus, and by the possession of a short stubby keel. 

Genus PROGONOCYTHERE Sylvester-Bradley 1948 

Progonocythere acuminata sp. nov. 
(PL II, figs. 5-10 ; PL 12, figs. 1-4) 

Diagnosis. Progonocythere with posteriorly acuminate carapace. Ventro-lateral 
margin convex with knife-edge keel developed in some specimens. Lateral surface 
smooth, though may possess faint transverse furrows medially. Low eye swelling 
at anterior cardinal angle. Ventral surface with longitudinal ridges in region of 
ventro-lateral overhang. Anterior with well developed border. 

Holotype. I0.1783, bed 7, Hundale Point, Cloughton. 

Paratypes. Io. 1784-91, bed 7, Hundale Point ; bed 6, Cayton Bay ; bed 7, 
Bogmire Gill ; top bed 7, Ravenscar and bed 10, Hawsker. 

Description . Carapace elongate , tapering strongly towards the sharply acuminate 
posterior. Sexual dimorphism not apparent. The ventro-lateral border of the 
carapace is extended below the ventral surface and generally possesses a thin, knife- 
edge keel, developed from one of the longitudinal ridges which extend along the 
undersurface of the ventro-lateral part of the carapace, remainder of ventral surface 
smooth. The ventro-lateral margin of the carapace sweeps obliquely upwards just 
behind valve middle. Shell surface smooth, although weak transverse furrows may 
be observed in some specimens in the mid-lateral area. Normal pore canals often well 
developed, large and circular, widely scattered over the carapace. Greatest length 



MIDDLE JURASSIC OSTRACODA 



115 



through mid-point ; greatest height in the anterior third ; greatest width at or just 
behind middle. A shallow, indistinct sulcus, medially situated marks the position 
of the adductor muscle scars. Anterior broadly rounded with a distinct marginal 
border ; posterior narrow, acuminate with a short, concave postero-dorsal slope and 
a convex postero- ventral slope. Ventral margin broadly concave. Dorsal margin 
slightly convex in the right valve slightly concave medially in the left. Cardinal 
angles prominent. Below the anterior cardinal angle an oblique swelling, separated 
off from the convex part of the carapace below by an oblique groove, is suggestive of 
an eye swelling. Left valve larger than the right which it overlaps along the ventral 
margin and overreaches along the dorsal margin and around the anterior. The left 
valve may also overreach the right along the postero-dorsal slope but not at the 
extreme posterior. Hinge poorly seen in a single individual (left valve) where the 
terminal sockets are separated by a median groove which can be made out as dentate. 
Accommodation groove elongate, deep. Duplicature appears to be of moderate 
width, but imperfectly seen. Other internal details not observed. 



Dimensions 

Holotype, I0.1783, carapace (PI. 12, figs. 1-4), length o-8o mm. ; height 0-49 mm. ; 
width 0-38 mm. 

I0.1784, left valve, length 071 mm. ; height 0-41 mm. I0.1786, left valve (PL 
II, figs. 7, 8), length o-6o mm. ; height 0-36 mm. I0.1787, carapace (PL 11, figs. 9, 
10), length 0-69 mm. ; height 0-41 mm. ; width 0-34 mm. I0.1789, carapace (PL 
II, figs. 5, 6), length 073 mm. ; height 0-41 mm. ; width 0-36 mm. I0.1791, right 
valve, length 071 mm. ; height 0-36 mm. 

Remarks. The similarity of P. acuminata to the male dimorph of Malzia uni- 
carinata has already been discussed and need not be gone into again. The male 
dimorph of Progonocythere yonsnabensis sp. nov. also bears some resemblance to the 





Figs. 16-17. Dorsal and left views, female carapace, Progonocythere vonsnabensis sp. nov. 

Holotype, lo. 1792, approx. X95. 



ii6 



R. H. BATE 



present species although it can be readily distinguished by its much smaller size and 
by the distinct median sulcus not present to such an extent here. 

Progonocythere yonsnahensis sp. nov. 
(PL 12, figs. 5-14 ; PI. 13, figs. 1-4 ; Text-figs. 16-19) 

Diagnosis. Small progonocytherid with distinct median sulcus. Eye swelling 
prominent. Ventro-lateral margin extended into ventral keel. Shell surface 
punctate. 

HoLOTYPE. I0.1792, bed 5, Cayton Bay section. 

Paratypes. Io. 1793-96, horizon and locality as above. 

Description. Carapace small, subquadrate ; elongate in the male dimorph. 
Greatest length through mid-point ; greatest height in the anterior third ; greatest 
width in the posterior third. As seen in dorsal view a shallow sulcus divides the cara- 
pace into an anterior and a posterior half. This sulcus is more strongly developed 
in the female. Ventro-lateral margin strongly convex, extended ventrally into a 
prominent keel. In a single male specimen there is a second, short keel dorsal to the 
first and terminating in a small node. Almost directly above this node a second node 
is situated dorso-medially on each valve of the carapace posterior to the median 
sulcus. In all other respects this specimen is identical to the others. Dorsal margin 
strongly convex in the left valve, slightly convex in the right. Anterior rounded 
with a short, obliquely convex antero-dorsal slope. Posterior triangular : postero- 
dorsal slope concave ; postero-ventral slope convex. Ventral margin medially 
incurved. Cardinal angles prominent, the posterior angle being the more strongly 
angled of the two, anterior angle tending to be broadly rounded. A prominent oval 





Figs. 18-19. Left and dorsal views, male carapace, Progonocythere yonsnahensis sp. nov., 
showing development of second keel and tubercles. Paratype, I0.1795, approx. X95. 



MIDDLE JURASSIC OSTRACODA 117 

eye swelling is situated below the anterior cardinal angle. Anterior and posterior 
marginal borders distinct. Shell surface punctate. Left valve larger than the right 
which it overlaps mid-ventrally and overreaches postero-ventrally, postero-dorsally 
and antero-dorsally. Mid-dorsally the left valve is strongly projected above the 
right. There is little or no overreach around the anterior margin. Internal charac- 
ters not observed for this species. 

Dimensions 

Holotype, I0.1792, female carapace (PI. 12, figs. 5-8 ; Text-figs. 16-17), length 
0-54 mm. ; height 0-32 mm. ; width 0-30 mm. 

I0.1793, male carapace (PI. 12, figs. 11-14), length 0-57 mm. ; height 0-34 mm. ; 
width 0-31 mm. 10.1794, female carapace (PL 12, figs. 9, 10), length o-6o mm. ; 
height 0-36 mm. ; width 0-35 mm. I0.1795, male carapace (PI. 13, figs. 1-4 ; Text- 
figs. 18-19), length 0-63 mm. ; height 0-36 mm. ; width 0-35 mm. 

Remarks. The single specimen (I0.1795) mentioned in the description, possessing 
two ventro-lateral keels and two lateral nodes is considered to be simply an extreme 
variant of the present species. Possibly this morphological variation may have been 
brought about by changes in the salinity of the water. Morphologically the posses- 
sion of two lateral keels results in this specimen having some resemblance to Malzia 
bicarinata, although it can be distinguished by the possession of a prominent median 
sulcus and a very much smaller adult size. 

The species Progonocy there yonsnabensis more closely resembles Progonocythere 
cristata Bate (1963 : 191, pi. 4, figs. 5-15 ; pi. 5, figs. 1-6) than any other ostracod. 
It is, however more strongly sulcate, possesses a more prominent eye swelling and is 
very much smaller. 

Subfamily PLEUROCYTHERINAE Mandelstam i960 

Genus PLEUROCYTHERE Triebel 1951 

Pleurocythere sp. 

(PI- 13, fig- 5) 
Remarks. A single left valve belonging to the genus Pleurocythere but not 
readily assignable to any known species is here recorded from the Gristhorpe Bay 
section, bed 4. The ornamentation of the valve consists of a longitudinal ventro- 
lateral ridge, an oblique median ridge and a short diagonal ridge which is situated 
below the anterior cardinal angle. This short ridge V's downwards, connected by a 
short ridge at the apex of the V to the median ridge. The dorsal end of the diagonal 
ridge terminates in an eye swelling. Shell surface between the ridges strongly 
reticulate. 

Dimensions. I0.1836, left valve (PI. 13, fig. 5), length o-8o mm. ; height 0-35 mm. 

Family CYTHERIDEIDAE Sars 1925 

Subfamily CYTHERIDEINAE Sars 1925 

Genus VERNONIELLA Oerth 1957 



ii8 R. H. BATE 

Remarks. The genus as diagnosed by Oertli (1957 : 659) possesses either a hemi- 
merodont or antimerodont type hinge, and is considered to be without any strong 
ornamentation. Two species are here placed in Vernoniella : V. bajociana sp. nov., 
a smooth form possessing an antimerodont hinge and the strongly ornamented V. ? 
caytonensis sp. nov. There is, however, some uncertainty concerning the generic 
designation of the last named species. 

Vernoniella bajociana sp. nov. 
(PI. 13, figs. 6-11 ; PI. 14, figs. 1-13) 

Diagnosis. Vernoniella with subquadrate carapace, elongate in male dimorphs. 
Shell surface finely punctate. Anterior and posterior margins with narrow marginal 
borders. Hinge antimerodont. Radial pore canals long, straight, approximately 
lo-ii anteriorly, 4-5 posteriorly. 

HoLOTYPE. I0.1807, bed 23, Hundale Point, Cloughton. 

Paratypes. Io. 1808-30 and I0.2988, bed 5, Cayton Bay ; beds 7, 8, 23 and 25, 
Hundale Point ; bed 12, Ravenscar and beds 9 and 10, Hawsker. 

Description. Carapace subquadrate to subtriangular in the female dimorph, 
elongate in the male. Shell surface punctate. Ventral surface may possess weak 
longitudinal ridges. In the dorso-median part of the carapace the valve is slightly 
constricted and here may exhibit weak, transverse furrowing. Greatest length of 
carapace through mid-point ; greatest height in the anterior third ; greatest width 
in the posterior third, although there is here only a slight increase over the width in 
the anterior third. Carapace appears compressed in dorsal view, almost parallel-sided. 
Dorsal margin medially concave in the left valve (both dimorphs), convex in the right. 
Cardinal angles in both valves prominent : anterior angle broadly rounded, posterior 
angle sharply angled and situated at extreme posterior of the carapace. Ventral 
margin medially incurved. Ventral surface overhung slightly by the convex ventro- 
lateral margin. The incurving of the ventral surface produces, particularly in the 
male, an enlarged anterior portion of the carapace as seen in lateral view. Anterior 
broadly rounded with marginal border which is separated off from the convex lateral 
part of the carapace by a marginal groove. Posterior broadly triangular with a 
steeply angled, straight or slightly convex postero-dorsal slope and a convex postero- 
ventral slope. Left valve larger than the right which it overlaps along the ventral 
and postero-ventral margins and overreaches around the anterior margin and along 
the postero-dorsal slope. Mid-do rsally the convex dorsal margin of the right valve 
projects above the left. Hinge antimerodont : left valve with terminal loculate 
sockets and a strong denticulate median bar. No accommodation groove. Right 
valve with 6 terminal teeth, dorsally bifid and an elongate, finely locellate median 
groove. Mtiscle scars (Type B?) with a sub vertical row of 4 oval adductor scars, a 
rounded antero-ventral mandibular scar which as seen in one individual is composed 
of a rosette of several small scars and an antero-dorsal antennal scar which is clover- 
leaf in shape. The antennal scar appears to be formed by the fusion of at least three 



MIDDLE JURASSIC OSTRACODA 119 

scars which together form a clover-leaf pattern, or may appear rounded. Between 
the antennal scar and the dorsal adductor scar there is a large depression which is 
the mandibular support spot. This muscle scar arrangement is tentatively placed 
as Type B. Inner margin and line of concrescence coincide, the duplicature being of 
moderate width. Radial pore canals straight, approximately lo-ii anteriorly and 
4-5 posteriorly. Outside the selvage in the right valve ^. flange is developed around 
the anterior margin, extending back along the ventral margin, where it expands 
opposite the ventral incurvature into a broad " lip ". 

Dimensions 

Holotype, I0.1807, female carapace (PI. 13, figs. 6-9), length 0-67 mm. ; height 
0-38 mm. ; width 0-30 mm. 

lo. 1808, female carapace (PI. 13, figs. 10, 11), length o-66 mm. ; height 0-37 mm. 
width 0-26 mm. I0.1812, female left valve, length o-6g mm. ; height 0-44 mm. 
I0.1814, male carapace (PI. 14, figs. 1-4), length 0-84 mm. ; height 0-43 mm. ; width 
0-29 mm. I0.1816, female carapace, length o-6i mm. ; height 0-34 mm. ; width 
0-28 mm. I0.1818, female left valve (PL 14, figs. 5-9), length 0-48 mm. ; height 
0-31 mm. I0.1819, female right valve (PL 14, figs. 10, 12, 13), length 0-65 mm. ; 
height 0-37 mm. I0.1821, male carapace, length o-68 mm. ; height 0-37 mm. ; 
width 0-28. 

Remarks. Vernoniella bajociana is similar in general external appearance to 
V . seqtiana Oertli (1957 : 659, pi. 3, figs. 70-85) but has an antimerodont instead of a 
hemimerodont hinge, and also lacks the accommodation groove which is present in 
V. sequana. V. bajociana has a distinct marginal border, a feature not present in 
Oertli's species. 

Vernoniella ? caytonensis sp. nov. 
(PL 15, figs. 1-9) 

Diagnosis. Vernoniella ? with elongate, posteriorly acuminate carapace. Shell 
surface strongly ornamented by 4-5 longitudinal ridges arranged in broad, inverted 
V. Two to three obliquely transverse ridges occur anterior to these. Anterior and 
posterior with distinct marginal borders. Normal pore canals prominent. 

Holotype. I0.1855, bed 6, Cayton Bay section. 

Paratypes. Io. 1856-69, bed 4, Gristhorpe Bay ; bed 6 Cayton Bay and bed 11, 
Hawsker. 

Description. Carapace elongate tapering towards the narrowly rounded pos- 
terior. Greatest length through mid-point ; greatest height in the anterior third ; 
greatest width in the posterior third. In dorsal view the carapace is slightly con- 
stricted medially. Anterior broadly rounded ; posterior narrowly rounded with 
steeply angled, convex postero-dorsal slope and convex postero-ventral slope. 
Dorsal margin in the right valve medially convex, in the left valve thickened and 
medially concave. Cardinal angles prominent ; the anterior angle being broadly 



I20 R. H. BATE 

rounded, posterior angle more acute, situated at extreme posterior of carapace. 
Anterior and posterior with distinct marginal borders. Ventral margin medially- 
incurved. Shell surface strongly ornamented by 4-5 longitudinal ridges arranged 
in a broad inverted V, the lowermost being almost straight. These ridges are 
situated on the strongly convex part of the carapace which is cut off posteriorly by 
the posterior border and anteriorly by an oblique groove running just behind and 
below the anterior cardinal angle. In front of this groove there are 2-3 obliquely 
transverse ridges which antero-ventrally bend back to extend along the ventro- 
lateral and ventral surfaces where weak longitudinal ridges may be made out. 
Normal pore canals distinctly se;n along the ventral surface where they are large and 
circular, elsewhere tending to be masked by lateral ornamentation. Left valve 
larger than the right which it overlaps along the ventral margin and overreaches in 
the region of the antero-dorsal and postero-dorsal slope. Hinge incompletely seen : 
right valve with the terminal elements consisting of approximately 6 dorsally bifid 
teeth. Median groove obscured by matrix. Left valve hinge not seen. Inner 
margin and line of concrescence appear to coincide. Radial pore canals long, straight 
and widely spaced, approximately 8 anteriorly and 3-4 posteriorly. Below the 
ventral incurvature in the right valve an elongate " lip " is developed. 

Dimensions 

Holotype, I0.1855, carapace (PL 15, figs. 1-4), length 0-63 mm. ; height 0-33 mm. ; 
width 0-30 mm. 

I0.1857, right valve (PL 15, figs. 8, 9), length o-66 mm. ; height 0-33 mm. I0.1858, 
carapace (PL 15, figs. 5-7), length 0-66 mm. ; height 0-37 mm. ; width 0-32 mm. 

Remarks. In outline this species approaches Vernoniella and as a result has been 
placed tentatively in that genus. It is, however, a strongly ornamented form whose 
hinge and muscle scars are as yet unknown. 

Vernoniella ? caytonensis resembles the male dimorph of Eocytheridea carinata Bate 
(1964 : 18, pi. 5, figs. 5-8) primarily because of the similarity of ornament although it 
is easily distinguishable by the more prominent posterior cardinal angle, the not so 
strongly tapered carapace, and by the more strongly pronounced marginal borders. 

Genus LJUBIMOVELLA Malz 1961 
Ljubimovella piriformis Malz 

(PL 15, figs. 10-13 ; PI- 16, figs. 1, 2) 

1949. Ostracod 96 Brand : 337, pi. 10 (fauna 2), fig. 4, pi. 14. 

1961. Ljubimovella piriformis Malz : 165, pi. 2, figs. 15-25 ; text-figs. 2-3. 

1962. Ljubimovella piriformis Malz : Brand & Fahrion : 134, pi. 20, fig. 33 ; table 9. 

Material. Twenty-three specimens examined from the Grey Limestone Series, 
of which the following have been registered in the British Museum collections : 
lo. 2103-08. 



MIDDLE JURASSIC OSTRACODA 121 

Description. Carapace piriform, very much enlarged in the anterior third, in 
which region it attains maximum height. Greatest length below mid-point ; greatest 
width at, or in front of valve centre. A strong, downwardly projected spine is 
developed at the postero-ventral angle whilst a much shorter spine may also be 
present on the lower half of the anterior margin. Shell surface smooth except for the 
ventral surface where a number of fine longitudinal striae occur. Normal pore canals 
circular and widely scattered over the surface. Dorsal margin slightly concave 
medially ; cardinal angles broadly rounded, although the posterior angle may be 
acute. Ventral margin with a strong median incurvature. Anterior broadly 
rounded ; posterior obliquely rounded with a relatively long, convex postero-dorsal 
slope and a short, almost vertical but slightly convex postero-ventral slope which 
terminates in the prominent posterior spine. Left valve larger than the right which 
it overlaps along the ventral margin. Along the antero-dorsal and postero-dorsal 
slopes the left valve tends to overreach the right, whilst mid-dorsally the right valve 
overreaches the left. Hinge hemimerodont : left valve without an accommodation 
groove ; right valve with smooth median groove and terminal dentate elements. 
Muscle scars not clearly seen. Inner margin and line of concrescence coincide. 
Radial pore canals short and widest at their base, 5 in number antero-ventrally and 
5 postero-ventrally, of which the lowermost passes down through the centre of the 
posterior spine. A well developed flange extends around the anterior margin and 
along the ventral margin in the right valve, not observed in the left valve probably 
because of the preservation of the material. 

Dimensions 

I0.2103, carapace, length 0-69 mm. ; height 0-34 mm. ; width 0-31 mm. I0.2106, 
carapace (PL 16, figs, i, 2), length 0-75 mm. ; height 0-40 mm. ; width 0-32 mm. 
I0.2107, juv, carapace (PI. 15, figs. 10, 11), length 0-44 mm. ; height 0-25 mm. ; 
width 0-22 mm. I0.2108, right valve (PI. 15, figs. 12, 13), length 072 mm. ; height 
0-37 mm. 

Remarks. The single juvenile specimen observed of this species exhibits a much 
enlarged anterior half of the carapace. It would appear that in subsequent instars 
of the species there is a proportionately greater increase in the posterior part of the 
carapace, a process almost certainly related to the development of the reproductive 
organs as the animal reaches maturity. 

Family SGHULERIDEIDAE Mandelstam 1959 

Subfamily SCHULERIDEINAE Mandelstam 1959 

Genus MESOCYTHERIDEA nov. 

Diagnosis. Schulerideinae with oval/subquadrate carapace. Central part of 
valve strongly cut off from remainder of carapace by oblique groove below the anterior 
and posterior cardinal angles, particularly well developed in right valve. Dorso- 
median part of right valve strongly projected above dorsal margin. Hinge anti- 

GEOL. II, 3 9 



122 R. H. BATE 

merodont with anterior socket of left valve cutting back into median bar. Radial 
pore canals long, slightly curved, lo anteriorly, 4 posteriorly. Inner margin and line 
of concrescence coincident. Muscle scars (Type C) as for family. Left valve larger 
than right. 

Type species. Mesocytheridea howardianensis sp. nov. 

Remarks. The genus (feminine) is close to Eocytheridea Bate {igS^a- : 35) from 
which it has probably evolved. Mesocytheridea is distinguishable from Eocytheridea 
by the presence of a strongly antimerodont hinge and a slight reduction in the num- 
ber of anterior radial pore canals (10 as against 14) which tend to be straighter. The 
species Eocytheridea carinata Bate (1964 : 18, pi. 4, figs. 6-11 ; pi. 5, figs. 1-8) is very 
close to this genus but has a hemimerodont hinge and the radial pore canals are 
slightly more curved. 

Mesocytheridea howardianensis sp. nov. 
(PI. 16, figs. 3-11 ; PI. 17, figs. 1-3 ; Text-figs. 20, 21) 

Diagnosis. Mesocytheridea with oval/subquadrate carapace, convex in dorsal 
view. Ornamentation consists of some 5-6 low, broad ridges arranged in an inverted 
V. The lowermost, ventro-lateral ridges are longitudinal, thereby forming base to V. 
Ornamentation never strong, may be almost completely lacking, when shell appears 
smooth. Shell surface with prominent, circular, normal pore canals. Hinge, 
muscle scars, radial pore canals as for genus. Left valve larger than right. Species 
dimorphic. 

Holotype. I0.1870, bed 2, Stonecliff Wood, locality SE/740675. 

Paratypes. I0.1871-81, horizon and locality as above and i ft. 6 in. and 3 ft. S in. 
from base bed 8, Stonecliff Wood, locality SE/743676. 




Fig. 20. Right view, female carapace, Mesocytheridea howardianensis sp. nov. Holotype 

I0.1870, approx. XQS- 



Id. 1 870, approx. X93. 

Description. Carapace oval/subquadrate, the male dimorphs being more elongate 
in outline. Greatest length through mid-point ; greatest height in the anterior 
third ; greatest width in the posterior third. Dorsal margin medially concave in the 
left valve with well roimded cardinal angles. In the male dimorph the dorso-median 



MIDDLE JURASSIC OSTR A CODA 123 

part of the left valve tends to project slightly above the dorsal margin, but not so 
strongly as in the right valve. Cardinal angles in the right valve more acute than in 
the left. Ventral margin medially incurved, convex in the posterior half where the 
carapace appears to be noticeably swollen. Anterior rounded with oblique, straight 
or slightly convex antero-dorsal slope. Posterior narrowly rounded with convex 
postero-dorsal and postero-ventral slopes. Narrow, convex, anterior and posterior 
marginal borders are separated from the lateral part of the carapace by a marginal 
groove. Dorsal margin of the left valve noticeably thickened. Shell surface with 
something like 5-6 low, rather poorly developed ridges which are arranged in the 
form of an inverted V, the apex of which reaches the dorsal margin, more noticeably 
developed in the right valve. Several of the lower, ventro-lateral ridges are straight 
and form a base to the inverted V. A strong diagonal groove separates the central 
part of the valve from the terminal parts, particularly in the right valve. The an- 
terior groove extends diagonally below the anterior cardinal angle whilst the posterior 
groove extends below the posterior angle. A transverse ridge extends from the 
anterior cardinal angle towards the antero- ventral angle. Remainder of shell 
surface punctate, although a few very minor ridges may connect some of the antero- 
dorsal ridges. The degree with which the ornamentation is developed varies with 
each individual, but is often so poorly represented that the specimen appears smooth. 



Fig. 21. Internal view, right valve showing hinge, Mesocytheridea howardianensis sp. nov. 

Paratype, lo. 1879, approx. X95. 

In all cases the normal pore canals which are large and circular, are prominently 
displayed over the shell surface. Left valve larger than the right which it overlaps 
along the ventral margin and overreaches practically everywhere else apart from mid- 
dorsally where the two valves are drawn apart. Hinge antimerodont : left valve 
with elongate terminal loculate sockets separated by a strong, denticulate median 
bar. The anterior socket, and to some extent the posterior socket also, tends to 
cut back into the median bar. There is virtually no accommodation groove. In the 
right valve the median groove is quite strongly locellate, whilst terminally there are 
5 anterior and 7 posterior teeth, all dorsally bifid and well developed. Muscle scars 
(Type C) consist of a slightly crescentic row of 4 oval adductor scars with an antero- 
median, round, antennal scar. No mandibular scar observed. Inner margin and 

GEOL. II, 3 9§ 



124 R. H. BATE 

line of concrescence coincide, duplicature being quite broad. Radial pore canals long 
and widely spaced, some straight others slightly curved ; lo anteriorly and 4 pos- 
teriorly. 

Dimensions 

Holotype, I0.1870, female carapace (PI. 16, figs. 3-6 ; Text-fig. 20), length 073 
mm. ; height 0-43 mm. ; width 0-38 mm. 

I0.1871, female right valve (PI. 16, fig. 9, PI. 17, figs. 2, 3), length 071 mm. ; 
height 0-37 mm. I0.1872, male left valve (PL 17, fig. i), length 0-93 mm. ; height 
0'47 mm. I0.1875, female left valve (PI. 16, fig. 11), length 076 mm. ; height 0-44 
mm. I0.1877, male right valve, length 071 mm. ; height 0-36 mm. I0.1879, male 
right valve (PI. 16, figs. 8, 10), length 078 mm. ; height 0-41 mm. I0.1881, female 
left valve (PI. 16, fig. 7), length 071 mm. ; height 0-41 mm. 

Remarks. This is a much larger species than Eocytheridea carinata with which it 
bears some resemblance, although in M. howardianensis the ornamentation tends to 
be much weaker and the presence of a strong antimerodont hinge and a slight reduc- 
tion in the number of radial pore canals help to distinguish the two species. The 
majority of the specimens examined come from the uppermost beds of the Grey 
Limestone Series as exposed along the western shore-line. However a population 
belonging to this species has been found at the base of the Series in the Bloody Beck 
section. The specimens here are generally smaller, no males having been found and 
are very poorly ornamented. They are not, however, considered to be sufficiently 
distinct as to be separated specifically. 

Genus PRAESCHULERIDEA Bate 1963 

Praeschuleridea subtrigona (Jones & Sherborn 1888) 

Synonymy. See Bate (1964 : 22). 

Remarks. Two subspecies of P. subtrigona have been recognized ; P. subtrigona 
subtrigona having a size range of up to 0-56 mm. in the female dimorph and 0-58 mm. 
in the male ; and P. subtrigona magna where the range extends to 073 mm. for the 
female and 0-83 mm. for the male. Within the Grey Limestone Series, however, 
there is a third subspecies having a maximum size of about 0-68 mm. for the female 
and 077 mm.-o-82 mm. for the male. There are also a number of minor morphological 
details which help to distinguish this third subspecies. As all the specimens examined 
from the Grey Limestone Series fall into this intermediate range the subspecies 
Praeschuleridea subtrigona intermedia subsp. nov. has been erected to account for 
them. 

Praeschuleridea subtrigona intermedia subsp. nov. 
(PL 17, figs. 4-10 ; PL 18, figs. 1-9) 

Diagnosis. A subspecies of Praeschuleridea subtrigona in which adult female is 
of the order of o-68 mm. and male is 077 mm. to 0-82 mm. Carapace subtrigonal, 



MIDDLE JURASSIC OSTRACODA 125 

punctate. Posterior dorsal margin virtually straight, sloping strongly to posterior. 
Anterior dorsal margin long, obliquely convex. Posterior with steeply inclined, 
convex postero- ventral slope. Anterior with narrow marginal border ; posterior 
border poorly developed. 

HoLOTYPE. I0.1837, bed 7, Hundale Point, Cloughton. 

Paratypes. Io. 1838-54, bed 4, Gristhorpe Bay ; bed 12, White Nab, Scar- 
borough ; bed 7 & base bed 22, Hundale Point and bed 2 Stonecliff Wood, locality 

SE/740675. 

Description. Carapace subtrigonal in outline, more elongate in the male dimorph. 
Shell surface finely punctate. Normal pore canals fairly large, circular, evenly 
scattered over carapace. A very low eye swelling may be seen on the right valve, 
female dimorph and slightly more strongly developed in the right valve of the male, 
situated below the anterior cardinal angle, not observed in the left valve of either 
dimorph. Greatest length passes through mid-point ; greatest height and width 
approximately at centre of carapace. A narrow marginal border delimited along 
its inner side by a marginal groove extends around the anterior margin, only poorly 
developed around the posterior. Left valve larger than the right which it overlaps 
along the ventral margin and overreaches elsewhere around the carapace. Dorsal 
margin " umbonate " the highest point being just about at valve centre, the dorsal 
margin sloping steeply away from this point to the anterior and posterior. Anteriorly 
the dorsal margin is broadly convex and passes into the anterior margin without a 
break. Posteriorly the dorsal margin is steeply angled, straight or very slightly 
convex. Posterior cardinal angle more prominently developed than the anterior 
angle. Posterior rounded-triangular with a short postero-dorsal slope which is 
convex in the left valve and straight in the right. Postero- ventral slope longer, 
convex and tending to be obliquely angled away from the ventral margin. Anterior 
uniformly rounded. Ventral margin antero-medially incurved, medially convex. 
Hinge paleohemimerodont ; left valve with terminal loculate sockets and a short 
median bar (longer in the male dimorph) across which there is a narrow groove con- 
necting the terminal sockets. Accommodation groove broad and shelf-like. Right 
valve with strongly dentate terminal elements, not clearly seen in the present material. 
Muscle scars (Type C) with rounded anteromedian antennal scar. Inner margin and 
line of concrescence coincide ; duplicature of moderate width. Anterior radial pore 
canals slightly curved and in some specimens appearing to thicken slightly towards the 
outer termination ; 12-16 observed in the present material. 

Dimensions 

Holotype, I0.1837, female carapace (PL 17, figs. 4-6), length 0-64 mm. ; height 
0-40 mm. ; width 0-32 mm. 

I0.1838, male carapace, length 0-82 mm. ; height 0-45 mm. ; width 0-37 mm. 
I0.1839, male carapace (PL 18, figs. 1-3), length 0-77 mm. ; height 0-43 mm. ; width 
0-33 mm. I0.1840, female carapace (PL 17, figs. 7-10), length 0-62 mm. ; height 
0-41 mm. ; width 0-32 mm. I0.1843, female right valve (PL 18, fig. 9), length o-6o 



126 R. H. BATE 

mm. ; height 0-34 mm. I0.1846, female left valve (PL 18, figs. 6, 7), length o-6i mm. ; 
height 0-43 mm. 

Remarks. Apart from the variations in size range between the three subspecies 
of Praeschulerideasubtrigona there are also a number of additional characters by 
which the subspecies may be distinguished. The first of these characters concerns 
the angularity of the carapace which in the female dimorph of intermedia is very close 
to that of the type subspecies, the dorsal margin sloping strongly away from the 
region of greatest height. If anything, however, the posterior half of the dorsal 
margin is more elongate and not so steeply sloping as in subtrigona suhtrigona. The 
dorsal margin in magna appears more uniformly convex on either side of the region 
of greatest height, contrasting with the almost straight posterior part in the other two 
subspecies. Posteriorly there are slight differences in all three : triangular in 
subtrigona subtrigona ; rounded in magna and bluntly flattened in intermedia where 
the postero-ventral slope although convex appears to flatten out slightly on ap- 
proaching the extreme posterior. Actually the posterior margin of intermedia is 
closer to that of magna than to subtrigona subtrigona. The male dimorph of inter- 
media may be equivalent in size to that of magna — there however, it is the male of 
the last named species which shows the greater degree of angularity, being more 
noticeably " umbonate " than intermedia. 

The conditions pertaining in north-eastern England during Middle Jurassic times 
appear to have offered a number of environments inhabited by a subspecies of 
Praeschuleridea subtrigona each population being characterized by a variation in 
size range. Praeschuleridea subtrigona subtrigona inhabited the marine waters of the 
shallow oolitic sea which covered Lincolnshire during Bajocian times. The northern 
extension of this sea lapped against the Yorkshire delta and provided a changed 
environment in which the subspecies P. s. magna developed. Higher up in the 
succession, the Yorkshire delta was still influencing the sedimentation of the Bajocian 
and throughout the entire marine embayment which spread over N.E. Yorkshire 
the population that existed was of P. s. intermedia. 

Family CYTHERURIDAE Miiller 1894 
Genus EOCYTHEROPTERON Alexander 1933 

Eocytheropteron ' sp. 
(PI. 18, figs. 10-13 ; PL 19, figs. 1-4) 

Remarks. Four complete carapaces of a species externally resembling the genus 
Eocytheropteron have been obtained from beds high up in the Grey Limestone Series. 
Two male carapaces (lo.igio-ii) have been obtained from bed 8, Stonecliff Wood 
section whilst a single female carapace (I0.2102) has been obtained from bed 5 of the 
same section. From bed 11, Hawsker, a further female carapace (I0.1909) has been 
found. The carapace is elongate-oval in side view with the ventro-lateral margin 
strongly convex and overhanging the ventral surface, especially so in the female 
dimorph. A short caudal process is developed and the greatest length of the cara- 



MIDDLE JURASSIC OSTRACODA 



127 



pace extends through mid-point. Shell surface with a very faint reticulation. A 
shallow sulcus is present at about valve centre in the female, slightly anterior to this 
in the male. Left valve larger than the right. Internal details not known although 
some radial pore canals can be made out externally in the antero-ventral area where 
they appear to be straight and widely spaced. This species is close to Cytheropteron 
(Cytheropteron) punim Schmidt (1954 : 88, pi. 6, figs. 3-6 ; pi. 7, figs. 25-29) although 
somewhat smaller and may be further distinguished by the more elongate and pos- 
teriorly tapering carapace of the female dimorph which in C. (C.) purum is shorter in 
comparison. 

I0.1909, female carapace (PI. ig, figs. 1-4), length 0-47 mm. ; height 0-26 mm. ; 
width 0-26 mm. I0.1911, male carapace (PL 18, figs. 10-13), length 0-53 mm. ; 
height 0-26 mm. width 0-28 mm. 

Genus PARACYTHERIDEA Miiller 1894 

Paracytheridea} caytonensis sp. nov. 

(PL 19, figs. 5-16 ; Text-figs. 22, 23) 

Diagnosis. Paracytheridea ? with backwardly projected ala terminating in 
prominent node. A second node is situated dorso-medially on lateral surface of valve, 
just behind median sulcus. Shell surface reticulate. Anterior and posterior with 
flattened marginal borders. Left valve larger than right. Species dimorphic. 

HoLOTYPE. I0.2137, bed 6, Cayton Bay Section. 

Paratypes. Io. 2138-42, horizon and locality as above. 





Figs. 22-23. Dorsal and left views, female carapace, Paracytheridea ? caytonensis sp. nov. 

Holotype, I0.2137, approx. x 105. 

Description. Carapace subquadrate in side view with a prominent, backwardly 
projected ala at the tip of which is situated an oval node-like swelling. Above and 
shghtly in front of this swelling a much larger node is situated on the lateral surface 
of the valve. In front of this circular node a transverse median sulcus is present 



128 R. H. BATE 

which appears to curve under the lateral node. An oblique groove extending from 
the dorsal margin, below the anterior cardinal angle, extends down to the antero- 
ventral part of the valve where it turns back to extend along the dorsal side of the 
ventro-lateral ala. Shell surface rather coarsely reticulate, including the surface of 
the two nodes. An eye swelling is situated just below the anterior cardinal angle. 
Anteriorly and posteriorly the marginal borders are flattened, the convex lateral part 
of the carapace not extending right up to the margins. Ventral surface with approxi- 
mately 5 longitudinal ridges to each valve. Sexual dimorphism indicated by the 
presence of an elongate specimen considered to be the male. Greatest length of 
carapace passes through mid-point ; greatest height in the anterior third ; greatest 
width in the posterior third. Dorsal margin slightly concave in the left valve, convex 
in the right. Anterior margin rounded with oblique, convex, antero-dorsal slope. 
Posterior triangular, extended into a caudal process, especially in the female. 
Postero-dorsal slope strongly concave, postero-ventral slope strongly convex. 
Ventral margin medially incurved. Cardinal angles prominent ; anterior angle 
broadly rounded ; posterior angle more acute. Left valve larger than the right which 
it overlaps along the ventral margin and slightly at the cardinal angles. Elsewhere 
the left valve overreaches the right, apart from along the dorsal margin where the 
valves diverge. Internal characters not seen. 

Dimensions 

Holotype, I0.2137, female carapace (PI. 19, figs. 5-8 ; Text-figs. 22, 23), length 
0-45 mm. ; height 0-26 mm. ; width 0-25 mm. 

I0.2138, male carapace (PI. 19, figs. 13-16), length 0-54 mm. ; height 0-25 mm. ; 
width 0-23 mm. I0.2140, female carapace (PI. 19, figs. 9-12), length 0-43 mm. ; 
height 0-25 mm. ; width (broken) 0-22 mm. 

Remarks. Only six specimens of this species have been found and these all occur 
in the same bed and represent a single population. Owing to the lack of knowledge 
concerning the internal details of this ostracod the generic designation is given with a 
query, although on general external morphology there is good reason to place this 
species into Paracytheridea. Superficially there is some resemblance between 
P. ? caytonensis and Cytheropteron (Cytheropteron) bispinosum crassum Schmidt 
(1954 : 87, pi. 7, figs. 23-24) although in the last named ostracod the ventro-lateral 
alar projection is not backwardly directed as in P. ? caytonensis and also lacks the 
characteristic nodes of that species. 

Family PROTOCYTHERIDAE Ljubimova 1955 
Subfamily KIRTONELLINAE Bate 1963 
Genus SOUTHCAVEA Bate 1964 

Southcavea microcellulosa sp. nov. 
(PI. 20, figs. 1-13 ; PI. 21, figs. 1-4) 
Diagnosis. Southcavea with oval-subquadrate carapace and coarse reticulate 



MIDDLE JURASSIC OSTRACODA 129 

ornament. Pits produced by reticulate ornament 4-6 sided, internally strongly 
punctate. Hinge antimerodont. Species dimorphic. 
HoLOTYPE. I0.1882, bed 5, Stonecliff Wood section. 

Paratypes. Io. 1883-99, beds 3, 5 & 8, Stonecliff Wood and bed 11, Hawsker. 

Description. Carapace oval-subquadrate in outline, more elongate in the male 
dimorph. Posteriorly tapered in the female. Greatest length through mid-point ; 
greatest height in the anterior third ; greatest width just behind valve centre. 
Dorsal margin in the left valve slightly concave medially with broadly rounded 
cardinal angles, in the right valve slightly convex, cardinal angles somewhat more 
acute. Anterior broadly rounded ; posterior more narrowly rounded with convex 
postero-dorsal and postero -ventral slopes in the left valve, and a concave postero- 
dorsal and convex postero-ventral slope in the right. Ventral margin medially 
incurved. Ventro-lateral margin convex. Anterior without a marginal border, 
whilst posteriorly there is a very narrow, flattened border. Left valve larger than 
the right which it overlaps mid-ventrally, at the cardinal angles and along the 
postero-dorsal slope. Antero-ventrally, postero-ventrally and antero-dorsally the 
left valve overreaches the right. Shell surface strongly reticulate, the 4-6 sided pits 
produced being strongly punctate. This degree of punctation in many cases has 
resulted in the pits themselves being subdivided by secondary ridges — this develop- 
ment appears to be more characteristic of the ventro-lateral areas of the carapace. 
Ventral surface strongly ornamented by longitudinal ridges. Normal pore canals 
few in number and widely scattered over the carapace, although because of the 
ornamentation only clearly seen along the ventral surface. Hinge antimerodont : 
left valve with terminal sockets separated by a finely denticulate median bar. The 
accommodation groove is elongate and shelf-like. Right valve with 5 bifid, posterior 
teeth, exact number anteriorly not known but appears to be more than 5. Median 
groove elongate and finely locellate. Inner margin and line of concrescence coincide, 
duplicature of moderate width. Radial pore canals straight, simple and widely 
spaced, approximately 8 anteriorly, at least 4 posteriorly. Muscle scars (Type D?) : 
adductor scars an oblique row of 4 scars with an antero-ventral mandibular scar and 
an antero-dorsal antennal scar which may be heart-shaped, but not so definitely V- 
shaped as in other species of this genus. A narrow flange has been observed in the 
right valve extending around the anterior margin and along the ventral margin. 

Dimensions 

Holotype, I0.1882, male carapace (PI. 20, figs. 1-4), length 0-67 mm. ; height 0-36 
mm. ; width 0-38 mm. 

I0.1883, female carapace (PI. 20, figs. 5-8), length o-6o mm. ; height 0-36 mm. ; 
width 0-37 mm. lo. 1886, female right valve (PI. 20, figs, g, 10), length (broken) 
o-6o mm. ; height 0-34 mm. I0.1888, male carapace (PI. 21, figs. 1-4), length 077 
mm.; height 0-41 mm. ; width 0-46 mm. ; I0.1891, femalecarapace,lengtho-65mm. ; 
height 0-36 mm. ; width 0-36 mm. I0.1893, female carapace, length 0-54 mm. ; 
height 0-30 mm. ; width 0-31 mm. 



I30 R. H. BATE 

Remarks. Southcavea microcellulosa is a much more elongate ostracod than 
Southcavea reticulata Bate (1964 : 27, pi. 10, figs. 3-14 ; pi. 11, figs. 1-4) with which 
it might be confused, and also has a much finer ornamentation. S. microcellulosa 
appears to be virtually restricted to the sandy shore-line facies along the western 
outcrop of the Grey Limestone Series with the exception of a single female carapace 
found in bed 11, Hawsker. However, even at Hawsker in the east it is doubtful 
whether the northern coast-line was very far away. 

Genus SYSTENOCYTHERE Bate 1963 

Sy St enocy there ovata sp. nov. 

(PI. 21, figs. 5-12) 

Diagnosis. Systenocythere with ovoid carapace showing a strong reticulate 
ornamentation when well preserved, but generally only possessing ornamentation of 
longitudinal ridges on ventro-lateral and ventral surfaces. Internal characters as for 
genus. 

HoLOTYPE. I0.1900, bed 7, Hawsker. 

Paratypes. I0.1901-8, bed 4, Gristhorpe Bay ; bed 6, Cayton Bay ; beds 19 
and 22, Hundale Point, Cloughton ; beds 7 and 12, Hawsker and bed i, Yearsley 
Moor. 

Description. Carapace ovoid in side view, tapering posteriorly. Greatest length 
through mid-point ; greatest height in the anterior third ; greatest width in the 
posterior third. Dorsal margin convex in both valves ; cardinal angles rounded. 
Anterior rounded with oblique, convex, antero-dorsal slope. Posterior narrowly 
rounded with convex postero-dorsal and postero- ventral slopes in the left valve and 
concave postero-dorsal and convex postero-ventral slopes in the right valve. Ventral 
margin medially incurved. Ventro-lateral margin convex, overhanging the ventral 
surface in side view. Anterior and posterior with flattened marginal borders. Left 
valve larger than the right which it overlaps most strongly mid-ventrally, from which 
point, along the ventral margin the left valve progressively overreaches the right. 
The left valve also overreaches the right along the antero- and postero-dorsal slopes. 
Shell surface may be either strongly reticulate or only noticeably ornamented along 
the ventro-lateral and ventral surfaces with longitudinal ridges. Fairly large normal 
pore canals may be seen, depending on preservation, widely scattered over the 
carapace. Hinge merodont, not fully determinable. Right valve with 5 anterior 
teeth and possibly 5 posterior teeth ; median groove long and very narrow, it is 
difficult to state precisely whether this groove is smooth or locellate because of the 
preservation of the material. Hinge poorly preserved in the left valve : Accom- 
modation groove of moderate width situated above the median bar which cannot be 
identified as being either smooth or denticulate. Muscle scars (Type D) consist of a 
subvertical row of 4 oval adductor scars with an oval anteroventral mandibular scar 
and a V-shaped antero-dorsal antennal scar. Inner margin and line of concrescence 
coincide, the duplicature being of moderate width. Approximately 8 anterior and 3 



MIDDLE JURASSIC OSTRACODA 



131 



posterior radial pore canals may be distinguished, these appear to be straight and 
widely spaced. A ndixrow flange widening opposite the ventral incurvature, extends 



GREY LIMESTONE SERIES 




Zone of 

Glyptocythere 

potita 

maximum 

thickness 

observed 

17 ft. II ins 


Zone of 

Glyptocythere 

scitula 

maximum 
thickness 
observed 
44 ft. 5 ins 


1 




Glyptocythere 




scitula 


i 




Fuhrbergiella(P) 




hornda hornda 






Praeschulendea 

subtrigona 

mtermedia 










Vernoniella 


^^^■i^ 




bajociana 






Monoceratina 




scarboroughensis 




1 1 II 


Progonocythere 




acuminata 






Ljubimovelta 
piriformis 






— i 


Cloughtonella 
rugosa 






Systenocythere 
ovata 






■ 


Paracypris 
bajociana 




^^m 


Eocytheropteron ? 
sp. 




1 


Pleurocythere sp. 






Caytonidea 




faveolata 




1 


Parac/theridea ? 
caytonensis 




^ 


Vernoniella ? 
caytonensis 




« 


Progonocythere 
yonsnabensis 






Mesocytheridea 










Southcavea 
microcellulosa 






Glyptocythere 
polita 






|o 


Glyptocythere 
costata 






1 


Malzia bicarinata 








Malzia unicannata 





Fig. 24. Range table of the Grey Limestone Series Ostracoda. 



132 R. H. BATE 

along the ventral margin of the right valve and possibly also extends around the 
anterior margin. A flange has not been observed in the left valve. 

Dimensions 

Holotype, I0.1900, carapace (PL 21, figs. 5-8), length 070 mm. ; height 0-44 mm. ; 
width 0-37 mm. 

I0.1901, left valve, length o-66 mm. ; height 0-39 mm. I0.1903, left valve (PI. 21, 
fig. 12), length o-6o mm. ; height 0-36 mm. I0.1904, right valve, length 0-48 mm. ; 
height 0-28 mm. I0.1907, right valve (PI. 21, figs. 9-11), length o-6o mm. ; height 
0-34 mm. 

Remarks. Systenocythere ovata is similar in outline to the female dimorph of 
S. exilofasciata Bate (1963 : 212, pi. 14, figs. 7-10 ; pi. 15, figs. 1-4), the type species, 
although it differs in not being so noticeably acuminate posteriorly and, when 
preservation permits, in being strongly reticulate. 

VI REFERENCES 
Arkell, W. J. 1933. The Jurassic System, in Great Britain, viii + 681 pp., 41 pis. Oxford. 
Bate, R. H. 1963. Middle Jurassic Ostracoda from North Lincolnshire. Bull. Brit. Mus. 

{Nat. Hist.) Geol., London, 8 : 173-219, pis. 1-15. 
• 1963a. Middle Jurassic Ostracoda from South Yorkshire. Bull. Brit. Mus. (Nat. Hist.) 

Geol., London, 9 : 19-46, pis. 1-13. 
■ 1964. Middle Jurassic Ostracoda from the Millepore Series, Yorkshire. Bull. Brit. Mus. 

{Nat. Hist.) Geol., London, 10 : 1-34, pis. 1-14. 
Black, M. 1928. " Washouts " in the Estuarine Series of Yorkshire. Geol. Mag. Lond., 65 : 

301-307. 
Brand, E. 1961. In Brand, E. & Malz, H. Ostracoden-Studien im Dogger, 3 : Drei neue 

Procytheridea-AvteTy und Ljubimovella n.g. aus dem NW-deutschen Bajocien. Senck. leth., 

Frankfurt a.M., 42 : 157-173, pis. i, 2. 
Brand, E. & Fahrion, H. 1962. Dogger NW — Deutschlands : 123-158, pis. 16-21. In 

Simon, W. & Bartenstein, H. (editors), Leitfossilien der Mikropaldontologie . viii + 432 

pp., 59 pis. Berlin. 
Brand, E. & Malz, H. 1962. Ostracoden-Studien im Dogger, 4 : Fuhrbergiella n.g. Senck. 

leth., Frankfurt a.M., 43 : 1-39, pis. 1-6. 
1962a. In Brand, E. & Fahrion, H. Dogger NW-Deutschlands : 123-159, pis. 16-21. 

In Simon, W. & Bartenstein, H. (editors), Leitfossilien der Mikropaldontologie. viii + 

432 pp., 59 pis. Berlin. 
Buckman, S. S. 1911. Appendi.x 1 — Ammonites from the Scarborough Limestone. Proc. 

Yorks. Geol. Soc, Leeds, 17 : 205-208. 
Dunbar, C. O. & Rodgers, J. 1958. Principles of Stratigraphy, viii + 356 pp., 123 text- 
figs. New York. 
Falcoln, N. L. & Kent, P. E. i960. Geological Results of Petroleum Exploration in Britain 

1945-1957. Mem. Geol. Soc. Lond., 2 : 1-56, pis. 1-5. 
Fox-Strangways, C. 1892. The Jurassic Rocks of Britain, 1, Yorkshire. Mem. Geol. Surv., 

London, ix -+- 551 pp. 
Hemingway, J. E. 1951. Report on a field meeting for 1949. Proc. Yorks. Geol. Soc, Leeds, 

28 : 118-122. 
Malz, H. 1959. Ostracoden-Studien im Dogger, 1 : Marslatoiirella n.g. Senck. leth., Frankfurt 

a.M., 40 : 19-23, 4 figs. 
1961. In Brand, E. & Malz, H. Ostracoden — Studien im Dogger, 3 : Drei neue Procy- 

theridea-Aiten und Ljubimovella n.g. aus dem NW-deutschen Bajocien. Senck. leth., 

Frankfurt a.M., 42 : 157-172, pis. i, 2. 



MIDDLE JURASSIC OSTRACODA 133 

Moore, R. C. 1961 (editor). Treatise on Invertebrate Palaeontology, Pt. Q., Arthropoda, 3. 

xxiii + 442 pp., 334 figs. Kansas. 
Oertli, H. J. 1957. Ostracodes du Jurassique Sup^rieur du Bassin de Paris (Sondage Vernon 

i). Rev. Inst, frang. Petrole, Paris, 12 : 647-695, pis. 1-7. 
Peterson, J. A. 1954. Jurassic Ostracoda from the " Lower Sundance " and Rierdon 

formation. Western Interior United States. /. Paleont., Tulsa, 28 : 153-176, pis. 17-19. 
Plumhoff, F. 1963. Die Ostracoden des Oberaalenium und tiefen Unterbajocium (Jura) 

des Gifhomer Troges, Nordwestdeutschland. Abh. Senckenb. natiirf. Ges., Frankfurt a.M., 

503 : i-ioo, pis. 1-12. 
Richardson, L. 1911. The Lower Oolitic Rocks of Yorkshire. Proc. Yorks. Geol. Soc, 

Leeds, 17 : 184-204. 
Schmidt, G. 1954. Stratigraphisch wichtige Ostracoden im " Kimeridge " und tiefsten 

" Portland " NW-Deutschland. Paldont. Z., Stuttgart, 28 : 81-101, pis. 5-8. 
Simon, W. & Bartenstein, H. (editors). Leitfossilien dex Mikropalaontologie. viii + 432 pp., 

59 pis. Berlin. J 

Sylvester-Bradley, P. C. 1948. Bathonian ostracoas from the Boueti Bed of Langton 

Herring, Dorset. Geol. Mag. Lond., 85 : 185-204, pis. 12-15. 
1956. The Structure, evolution and nomenclature of the ostracod hinge. Bull. Brit. Miis. 

{Nat. Hist.) Geol., London, 3 : 1-21, pis. 1-4. 



EXPLANATION OF PLATES 

All the specimens illustrated are now in the Department of Palaeontology, British 
Museum (Natural History) . All photographs, taken by the author, x 85 unless 
otherwise indicated. 



GEOL. tl, 3 



PLATE I 

Monoceratina scarboroughensis sp. nov. p. 99. 

Figs. 1-3, 12. Bed 22, Hundale Point. 

Figs. 4-11. Bed 5, Cayton Bay. 

Figs. 1-3. Right, left and ventral views, female carapace. Holotype, I0.1711. 

Fig. 4. Muscle scars, female carapace. Paratype, I0.1723, X310. 

Figs. 5-8. Right, left, dorsal and ventral views, female carapace. Parat^qse, I0.1723. 

Figs. 9-11. Left, dorsal and ventral views, male carapace. Paratype, 10.1721. 

Fig. 12. Internal view, female left valve. Paratype, I0.1720. 

Caytonidea faveolata gen. et sp. nov. p. 100. 
Specimen from bed 4, Gristhorpe Bay. 
Figs. 13, 14. Right, left, views, carapace. Paratype, I0.183.1. 



Bull.B.M. (N.H.) Geol. II, 3 



PLATE 1 





■» 






PLATE 2 

Caytonidea faveolata gen. et sp. nov. p. loo 

Figs. 1-4, 7-10. Bed 5, Cayton Bay. 

Figs. 5, 6. Bed 4, Gristhorpe Bay. 

Right, left, dorsal and ventral views, carapace. Holotype, I0.1831. 
Dorsal and ventral views, carapace. Paratype, I0.1834. 
Dorsal and lateral views of right valve hinge., Paratype, I0.1832, X95. 
Internal views of right valve fragment to show muscle scars and anterior radial 
pore canals. Paratype, I0.1833. 



Figs. 
Figs. 
Figs. 


I- 
5- 

7. 


-4- 
6. 
8. 


Figs. 


9, 


10. 



Bull. B.M. (N.H.) GeoL. 11, 3 



PLATE 2 




PLATE 3 

Cloughtonella rugosa gen. et sp. nov. p. 102 

Figs. 1-7, 11-13. Bed 22, Hundale Point, Cloughton. 

Figs. 8-10. Bed 10, Bloody Beck. 

Right, left, dorsal and ventral views, female carapace. Holotype, I0.2118. 
Ventral, right and left views, male carapace. Paratype, Io.2iig. 
Left and right views, female carapace. Paratype, I0.2135. 
Left view, showing large normal pore canals, male carapace. Paratype, I0.2134. 
Dorsal view of hinge, female left valve. Paratype, I0.2121. 
Figs. 12, 13. Right and left views, male carapace. Paratype, I0.2120. 



Figs, i- 


-4- 


FlGS. 5- 


"7- 


Figs. 8, 


9 


Fig. 10. 




Fig. II. 





Bull. B.M. [N.H.) Geol. 11, 3 



PLATE 3 




PLATE 4 

Fuhrbergiella (Praefuhrbergiella) horrida horrida Brand & Malz p. 104 

Figs. 1-3. Bed 5, Cayton Bay. 

Figs. 4, 5, 10. Bed 6, Cayton Bay. 

Figs. 6-9, 11, 12. Bed 23, Hundale Point, Cloughton. 

Figs. 1-3. Internal, external and dorsal views, female left valve, I0.2109. 
Figs. 4, 5. External and dorsal views, female right valve, I0.2116. 
Figs. 6-9. Dorsal, left, right and ventral views, male carapace, I0.2111. 
Fig. 10. External view, female right valve showing anterior and posterior radial pore canals, 
I0.2117. 

Figs, ii, 12. Dorsal and right views, female carapace, lo. 2110. 



Bull. B.M. (X.H.) Geol. 11, 3 



PLATE 4 




PLATE 5 

Glyptocythere costata sp. nov. p. io6 

Figs. 1-4, 7. Bed 10, Ravenscar. 

Figs. 5, 6. Bed 8, Ravenscar. 

Figs. 1-4. Right, left, dorsal and ventral views, female carapace. Holotype, I0.1775. 
Figs. 5, 6. Right and left views, male carapace. Paratype, I0.1782. 
Fig. 7. Female right valve. Paratype, I0.1776. 

Glyptocythere polita up. nov. p. 107 

Fig. 8. Bed 6, Bloody Beck. 

Fig. g. Bed 7, Ravenscar. 

Fig. id. Bed 7, Huudale Point, Cloughton. 

Fig. II. Bed 9, Ravenscar. 

Fig. 8. Right view, female carapace. Paratype, I0.1743. 

Fig. 9. Internal view, female left valve. Paratype, I0.1738, X70. 

Fig. 10. Right view, female carapace, showing normal pore canals. Parcitvpe, I0.1736, 
X70. 

Fig. II. Dorsal view of hinge, female left valve. Paratype, I0.1741. 



Bull. B.M. (N.H.) Geol. 11, 3 



PLATE 5 




PLATE 6 
Glyptocythere politasp. nov. p- 107 

Figs. 1-8. Bed 7, Hundale Point, Cloughton. 

Fig. 9. Bed 8, Hundale Point, Cloughton. 

Figs. 1-4. Right, left, dorsal and ventral views, female carapace. Holotype, I0.1724, x 70. 
Figs. 5-8. Right, left, dorsal and ventral views, male carapace. Paratype, I0.1725, x 70. 
Fig. 9. Muscle scars, female left valve. Paratype, I0.1737, Xi50- 



Bull. B.M. (N.H.) Geol. 11,3 



PLATE 6 




PLATE 7 

Glyptocythere scitula sp. nov, p. loS 

Figs, i, 7. Bed 10, Hawsker. 

Figs. 2-6, 8-10. Bed 12, Ravenscar. 

Figs, ii, 12. Bed 5, Cay ton Bay. 

Fig. 13. Bed 5, Gristhorpe Bay. 

Figs, i, 7. Internal and dorsal views, female right valve. Paratype, I0.1760. 

Fig. 2. Muscle scars. Note antero-ventral mandibular scar, which is a rosette of several, 
smaller scars. Broken female right valve. Paratype, lo. 1774, X 150. 

Figs. 3, 8. Muscle scars (X125) and anterior radial pore canals (xgo), female right valve. 
Paratype, lo. 1771. 

Figs. 4, 6, 10. Internal view, showing radial pore canals, and two dorsal views of hinge, 
female right valve. Paratype, I0.1768. 

Figs. 5, g. Dorsal views of hinge, female left valve. Paratype, I0.1769. 

Figs, ii, 12. External and internal views, male left valve. Paratype, 10.1731. 

Fig. 13. External view, male right vadve. Parats'pe, I0.1772. 



Btill. B.M. (N.H.) Geol. 11, 3 



PLATE 7 










10 



PLATE 8 

Glyptocythere scitula sp. nov. p. io8 

Figs. 1-5. Bed 5, Cayton Bay. 

Figs. 6-g. Bed 10, Hawsker. 

Figs. 1-4. Left, right, dorsal and ventral views, female carapace. 
Fig. 5. External view, female left valve. Paratype, I0.1754. 
Figs. 6-9. Left, right, dorsal and ventral views, female carapace. 



Holot\rpe, I0.1750. 
Paratv'pe, I0.1752. 



Bull. B.M. (N.H.) Geo!. 11, 3 



PLATE 




PLATE 9 
Glyptocythere scitula sp. nov. p. io8 
Specimen from bed lo, Hawsker. 
Figs. 1-4. Left, right, dorsal and ventral views, male carapace. Paratype, I0.1753. 

Malzia bicarinata gen. et sp. nov. p. 1 1 1 
Specimen from bed 9, Ravenscar. 
Figs. 5-8. Left, right, dorsal and ventral views. Holotype, I0.1797. 



Bull.B.M. {N.H.) Geol. 11, 3 




PLATE lo 

Malzia bicarinata gen. et sp. nov. p. iii. 

Fig. I. Bed 8, Ravenscar. 

Figs. 2, 3. Bed 7, Ravenscar. 

Fig. I. Left side, carapace. Paratype, I0.1798. 

Figs. 2, 3. Internal view and dorsal view ( x 100) left valve. Paratype, lo.ijgg. 

Malzia unicarinata gen. et sp. nov. p. 113. 

Figs. 4-8. Bed 9, Ravenscar. 

Figs. 9, 10. Bed 8, Ravenscar. 

Figs. 4-7. Left, right, dorsal and ventral views, female carapace. Holotype, I0.1801. 
Fig. 8. Right view of female carapace to show radial pore canals. Holotype, To. 1 801. 
Figs. 9, 10. Left and right views, female carapace. Paratype, I0.1802. 



Bnll.B.M. (X.H.) GeoL 11, 3 



PLATE 10 




PLATE I I 

Malzia unicarinata gen. et sp. nov. p. 113. 

Specimen from bed 10, Ravenscar. 

Figs. 1-4. Left, right, dorsal and ventral views, male carapace. Paratype, I0.1806. 

Progonocy there acuminata sp. nov. p. 114 

Figs. 5, 6. Bed 7, Ravenscar. 

Figs. 7-10. Bed 6, Cayton Bay. 

Figs. 5, 6. Left and right views, carapace. Paratype, I0.1789. 

Figs. 7, 8. External and internal views, left valve. Paratype, I0.1786. 

Figs, q, id. Right and left views, carapace. Parat^rpe, I0.1787. 



Bull.B.M. (X.H.) Geol. 11, 3 



PLATE 11 




PLATE I 2 
Progonocythere acuminata sp. nov. p. 114. 
Specimen from bed 7, Hundale Point, Cloughton. 
Figs. 1-4. Lett, right, dorsal and \entral views, carapace. Holotype, I0.1783. 

Progonocythere yonsnabensis sp. now p. iio. 

All specimens from bed 5, Cay ton Bay. 

Figs. 5-8. Left, right, dorsal and ventral views, female carapace. Holotype, 10.1792. 

Figs. <>, 10. Left and right views, female carapace. Paratype, I0.1794. 

Figs. 11-14. Left, right, dorsal and ventral views, male carapace. Parat},^e, 10.1793. 



Bull. B.M. (N.H.) Geol. 11, 3 



PLATE 12 




PLATE 1 3 

Progonocythere yonst\^ensis sp. nov. p. lib. 

Specimen from beds/ Cay ton Bay. 

Figs. 1-4. Left, right, dorsal and ventral views, male carapace, sho^ving development of two 
itenil l<cels and nodes. Paratjrpe, I0.1795. 

Pleurocythere sp. p. 117. 

Specimen from bed ^, Gristhorpe Bay. 

Fig. 5. External view, left valve, I0.1836. 

Vernoniella bajociana sp. nov. p. 118. 

Specimens from bed 2^, Hundale Point, Cloughton. 

Figs. 6-y. Left, right, dorsal and ventral views, female carapace. Holotype, I0.1807. 
Figs. 10, 11. Right and left views, female carapace. Parat^'pe, I0.1808. 



Bull.B.M. [N.H.) Geol. 11, 3 



PLATE 13 




PLATE 14 

Vertioniella bajociana sp. nov. p. 118. 

Figs. 1-4. Bed 7, Hundale Point, Cloughton. 

Figs. 5-10, 12, 13. Bed 12, Ravenscar. 

Fig. 1 1 . Bed g, Hawsker. 

Figs. 1-4. Right, left, dorsal and ventral views, male carapace. Paratype, I0.1814. 

Figs. 5-9. 5, anterior radial pore canals ( x 140) ; 6, dorsal view of hinge ; 7, internal view 
to show boring of shell by marine organism ; 8, lateral view of hinge ; 9, internal view to show 
radial pore canals. Female left valve. Paratype I0.1818. 

Figs. 10, 12, 13. Muscle scars ( x 180) showing composite antennal scar, dorsal view of hinge 
( X95) and internal view of male right valve. Paratype, I0.1819. 

Fig. II. Muscle scars, fragment of female right valve. Paratype, I0.1813, xiio. 



Bull. B.M. (N.H.) Geol. 11, 3 



PLATE 14 




PLATE 15 

Vernoniella caytonensis sp. nov. p. iig. 

All specimens from bed 6, Cayton Bay. 

Figs. 1-4. Left, right, dorsal and ventral views, carapace. Holotype, I0.1855. 
Figs. 5-7. Right, left and ventral views, carapace. Paratype, I0.1858. 
Figs. 8, 9. Dorsal and internal views, right valve. Paratype, I0.1857. 

Ljubimovella piriformis Ma.lz p. 120. 

Figs. 10, 11. Bed 22, Hundale Point, Cloughton. 

Figs. 12, 13. Bed 12, Ravenscar. 

Figs. 10, 11. Left and right sides, juvenile carapace. I0.2107. 
Figs. 12, 13. Internal and external views, right valve. I0.2108. 



B.M. (N.H.) Geol. 11, 3 



PLATE 15 




PLATE I 6 
LJubimovella piriformis Ma,\z. p. 120. 
Specimen from bed 7, Hawsker. 
Figs, i, 2, Left and right sides, adult carapace. I0.2106. 

Mesocytheridea howardianensis gen. et ^Tp. nov. p. 122. 

Figs. 3-6, 9, 11. Bed 2, Stonecliff Wood. 

Fig. 7. Bed 7, Bloody Beck. 

Figs. 8, 10. Bed 8, Stonecliff Wood. 

Figs. 3-6. Right, left, dorsal and ventral views, female carapace. Holotype, I0.1870. 
Fig. 7. Dorsal view hinge, female left valve. Paratype, I0.1881. 
Figs. 8, 10. Dorsal and internal views, male right valve. Paratvpe, I0.1879. 
Fig. 9. Dorsal view, female right valve. Paratype, I0.1871. 

Fig. II. Internal view, female left valve, showing anterior socket cutting back into the 
median bar. Paratv^pe, I0.1875. 



Bull. B.M. (N.H.) Geol. 11, 3 



PLATE 16 




PLATE 17 

Mesocyt her idea howardianensis gen. et sp. no\ . p. 122. 

Specimens from bed 2, Stonecliff Wood. 

Fig. I. Internal view, showing radial pore canals, male left valve. Paratype, lo. 1872. 
Figs. 2, 3. Internal views to show hinge and radial pore canals, female right valve. Paratype, 
I0.1871. 

Praeschuleridea subtrigona intermedia subsp. nov. p. 124. 

Figs. 4-6. Bed 7, Hundale Point, Cloughton. 

Figs. 7-10. Bed 2, Stonecliff Wood. 

Figs. 4-6. Left, right and dorsal views, female carapace. Holotype, I0.1837. 

Figs. 7-10. Ventral, dorsal, right and left views, female carapace. Paratype, I0.1840. 



BuU.B.M. {N.H.) Geol. 11, 3 



PLATE 17 




PLATE 1 8 

Praeschulerida subtrigona intermedia subsp. nov. p, 124. 

Figs. 1-3. Bed 7, Hundale Point, Cloughton. 

Figs. 4, 5, 8, 9. Bed 2, Stonecliff Wood. 

Figs. 6, 7. Bed 4, Gristhorpe Bay. 

Figs. 1-3. Left, right and ventral views, male carapace. Paratype, I0.1839. 
Fig. 4. Muscle scars from internal cast. Paratype, I0.1844, xi8o. 
Fig. 5. Anterior radial pore canals, right valve fragment. Paratype, I0.1841, X135. 
Figs. 6, 7. Dorsal and internal views to show hinge, female left valve. Paratype, I0.1846. 
Fig. 8. Right side to show normal and radial pore canals, female carapace. Paratype, lo. 
1842. 
Fig. 9. Dorsal view of hinge, female right valve. Paratype, I0.1843. 

Eocytheropteron ? sp. p. 126. 

Specimen from bed 8, Stonechff Wood. 
Figs. 10-13. Ventral, left, right and dorsal views, male carapace. I0.1911. 



BuU.B.M. {N.H.) Geol. 11, 3 



PLATE 18 




PLATE 19 

Eocytheropteron ? sp. p. 126. 

Specimen from bed 11, Hawsker. 

Figs. 1-4. Right, left, dorsal and ventral views, female carapace. I0.1909. 

Par acyt her idea ? cay tonensis sp. nov. p. 127 

All specimens from bed 6, Cayton Bay. 

Figs. 5-8. Dorsal, ventral, left and right views, female carapace. Holotype, I0.2137. 
Figs. 9-12. Right, left, dorsal and ventral views, female carapace. Paratype, I0.2140. 
Figs. 13-16. Right, left, dorsal and ventral views, male carapace. Paratype, I0.2138. 



Bull. B.M. (N.H.) Geol. 11, 3 



PLATE 19 




□n 




PLATE 20 

Southcavea microcellulosa sp. nov. p. 128. 

Figs. 1-8, 11-13. Bed 5, Stonecliff Wood. 

Figs. 9, 10. Bed 3, Stonecliff Wood. 

Figs. 1-4. Left, right, dorsal and ventral views, male carapace. Holotype, I0.1882. 

Figs. 5-8. Left, right, ventral and dorsal views, female carapace. Paratype, I0.1883. 

Figs. 9, 10. Dorsal view of hinge and muscle scars ( X120), female right valve. Paratype, 
I0.1886. 

Figs. 11-13. Dorsal view of hinge and internal views showing hinge and radial pore canals, 
female right valve. Paratype, I0.1885. 



Bull.BM. (N.H.) Geol. 11, 3 



PLATE 20 




PLATE 21 
Southcavea microcellulosa sp. nov. p. 128. 
Specimen from bed 8, Stonecliff Wood. 
Figs. 1-4. Right, left, dorsal and ventral views, male carapace. Paratype, I0.1888. 

Systenocy there ovata sp. nov. p. 130. 

Figs. 5-8. Bed 7, Hawsker. 

Figs. 9-11. Bed 6, Cayton Bay. 

Fig. 12. Bed 4, Cayton Bay. 

Figs. 5-8. Kight, left, dorsal and ventral views, carapace. Holotype, I0.1900. 

Figs. 9-1 i. External, internal and dorsal views, right valve. Paratype, I0.1907. 

Fig. 12. Muscle scars, note V-shaped antennal scar, left valve. Paratype, I0.1903, X240. 



Biill. B.M. {N.H.) Geol. 11, 3 



PLATE 21 




(' 




PRINTED IN GREAT BRITAIN 
BY ADLARD & SON LIMITED 
BARTHOLOMEW PRESS, DORKING 



.x^V' 



HUMAN SKELETAL MATERIAL %^^ ^^/ 

FROM CEYLON, WITH AN 

ANALYSIS OF THE ISLAND'S 

PREHISTORIC AND 

CONTEMPORARY POPULATIONS 



K. A. R. KENNEDY 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. ii No. 4 

LONDON: 1965 



HUMAN SKELETAL MATERIAL FROM CEYLON, 

WITH AN ANALYSIS OF THE ISLAND'S 

PREHISTORIC AND CONTEMPORARY 

POPULATIONS -^'^ 




BY 

KENNETH A. R. KENNEDY, Ph.D. 

(Department of Anthropology, Cornell University, Ithaca, New York) 



Pp. 135-213; 15 Plates; 9 Text-figures 
12 Tables 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

GEOLOGY Vol. 11 No. 4 

LONDON: 1965 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 4 of the Geological 
[Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 



Trustees of the British Museum (Natural History) 1965 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 8 December, 1965 Price £2 5s. 



HUMAN SKELETAL MATERIAL FROM CEYLON, 

WITH AN ANALYSIS OF THE ISLAND'S 

PREHISTORIC AND CONTEMPORARY 

POPULATIONS 

By K. A. R. KENNEDY 



CONTENTS 

Page 

I Introduction ......... 138 

The Site of Bellan Bandi Palassa ..... 138 

The Nature and Condition of the Specimens . . . . 141 

II Description .......... 145 

The Methodology of the Metrical Analysis .... 145 

Sex and Age Determination ...... 146 

The Metrical and Morphological Analysis of the Osseous Remains 150 

The Metrical and Morphological Analysis of the Dentition . 165 

The Biochemical Analysis . . . . . . . 173 

III Comparative Analysis ........ 183 

The Nature of the Comparative Data Used in the Determination 
of the Biological Affinities of the Balangodese with Other 

Populations . . ....... 183 

A Comparison with the Veddas . . . . . . 184 

IV Discussion .......... 201 

V Conclusions .......... 207 

VI Acknowledgements ........ 208 

VII References .......... 209 

SYNOPSIS 

The fossilized human skeletal remains from the prehistoric site of Bellan Bandi Palassa in 
Sabaragamuva Province, Ceylon, are analysed anthropometrically and biochemically. Results 
of uranium and radiocarbon assays indicate a relatively contemporary population living at a 
period around 114 b.c. ± 200 years, a date that confirms the archaeological evidence which 
ascribed the cultural associations of the site to the Bandarawelian, a regional variant of the 
Indian " Mesolithic " or Late Stone Age. Comparative anthropometric studies of these Balan- 
godese fossils with other hominid specimens, both fossil and living, reveal that their closest 
phenotypic affinities are with the Veddas of Ceylon. Many of the physical traits regarded by 
earlier investigators as distinguishing the Veddas from their Ceylonese and Indian neighbours 
and which have been the basis for separating the Veddas into racial sub-types are apparent in 
the fossil " pre-Vedda " remains from Bellan Bandi Palassa. This suggests a close genetic 
affinity between these Balangodese- Vedda phenotypes at the dawn of the historic period in 
Ceylon. The evidence from the ethnographic and prehistoric record for this region strongly 
supports the view of a continuous cultural tradition with local modifications extending from 
Late Stone Age times to the present, a situation which lends independent but supportive evidence 
for the postulation of Balangodese and Vedda phenotypic affinities. The association of con- 
temporary hill tribes or relict populations with the manufacturers of prehistoric lithic industries 
has hitherto been unsubstantiated in the Indian Sub-Continent, and the anthropological 
problems inherent in this line of research are discussed. 



GEOL. II, 4 



M 



138 HUMAN SKELETAL MATERIAL FROM CEYLON 

I INTRODUCTION 

The Site of Bellan Bandi Palassa 

The presence of palaeolithic artifacts in India and Pakistan testifies to the human 
occupation of the Sub-Continent during the Pleistocene, but the skeletal remains of 
the manufacturers of these stone implements have not been found. The most 
ancient human bones from this part of Asia have been recovered from Langhnaj in 
Northern Gujarat (Ehrhardt i960, 1963, Karve & Kurulkar 1945, Khan & Karve 
1946, Karve-Corvinus & Kennedy 1964, Kennedy 1964, Sankalia 1945, 1946, 1949, 
Sankalia & Karve 1944, 1945, 1949, Subbarao 1952, 1955 : 73-74, 1958 : 71-74, 
Zeuner 1950 : 44, 1951 : 7) and from the District of Mirzapur in Uttar Pradesh 
(Personal communication with Shri Radhakant Varma in May 1964, Deccan College, 
Poona) where the cultural associations are ascribed to the Indian Late Stone Age, 
or Mesolithic, period. For the Island of Ceylon the discovery of the makers of the 
lithic industries has long been awaited. The work of the Sarasins (1892-93, 1907) 
confirmed the claims of earlier investigators that the island possessed Stone Age 
tools, and from their data two problems originate : (i) To which of the prehistoric 
ages, as understood in terms of conventional typological classifications, can the 
Ceylonese stone artifacts be assigned : (2) What kinds of hominids manufactured 
these tools? 

" Until definite stratigraphic evidence, showing a sequence of distinct cultures 
with perhaps associated animal and skeletal remains, have been established at 
several sites, the only procedure seems to be to treat the implements as pro- 
visionally of one culture . . . Subdivisions into cultures and phases can be 
made when adequate evidence is discovered justifying it ... The Sarasins 
believed that the people who lived in the rock shelters and made stone imple- 
ments were ancestors of the Veddah, but without skeletal evidence there is 
nothing to support such a conclusion." (Noone & Noone 1940 : 20-21). 

It is with this latter problem that the present study is concerned, for the anthropo- 
metric analysis of the human remains from Bellan Bandi Palassa, Ceylon, indicates 
that the manufacturers of its Bandarawelian (Late Stone Age) industries bear striking 
phenotypic similarities to the surviving Vedda population of the island. 

The site of Bellan Bandi Palassa is situated at 6 degrees 31 minutes North longitude 
and 80 degrees 48^ minutes East latitude between the 400 and 300 foot contours in 
the Balangoda District of Sabaragamuva Province. Three and a half miles to the 
east of the site is the Pansadara Chena near Hath Kinda where the Valave Ganga 
ramifies into its seven channels. Traversing the site is an intermittent stream which 
joins the Pusalli Ara just before its junction with the Valave south of Pansadara 
(Text-fig. i). Within the palassa, or glade, of Bellan Bandi an expanse of flat 
crystalline limestone extends for about a quarter of a mile from the northwest to the 
southeast and has a width of 70 yards. At right angles to the stream bed and 
extending across the limestone exposure is a ridge of earth eroded by the stream to 
form a gap of some 60 feet across. This erosion has exposed on the left bank a 
kitchen midden some 30 feet wide and averaging 3 feet in height above the limestone. 



HUMAN SKELETAL MATERIAL FROM CEYLON 



139 



Some materials have been redeposited about 150 yards downstream from the midden. 
Erosion has produced along the left bank for a distance of about 100 yards a number 
of small rock shelters which have since collapsed leaving large limestone blocks which 
overlie the midden. The surface soil is primarily red sandy loam ranging in depth 
from 3 feet 6 inches to 4 feet 6 inches above the limestone outcrop. Within this 
soil cover were found human and animal bones, molluscan shells, chert and quartz 
implements and potsherds. 




Fig. I. The region of Bellan Bandi Palassa in the Balangoda District of Sabaragamuva 
Province, Ceylon. One inch to one and a half miles. 



The excavation was carried out from 24th June to 6th July, 1956, then resumed on 
9th September, 1956. A site survey had been conducted in March of that same 
year as a result of reports of fossil remains to be found in the forest near the Valave. 
Mr. Arthur Delgoda sent to Dr. P. E. P. Deraniyagala, Director of the National 
Museums, some fossilized fragments of human and animal bones, and on 12th March, 
1956 led a party of archaeologists under Dr. Deraniyagala to the site. Before the 
commencement of the excavation the site was pegged out into forty squares each 



140 



HUMAN SKELETAL MATERIAL FROM CEYLON 



6 feet by 6 feet. The north-south base Hne was represented by the plots Beta, 
Alpha, A, B, C, D, E, F, and the series beginning with Beta in the south end. The 
east-west base line was represented by the plots i, 2, 3, 4, and 5, the series beginning 
with 5 in the west end and adjacent to square Beta. Such a grid was superimposed 
on the site so that the areas of limestone exposure and the left bank of the stream 
were approximately equal in extent, the edge of the stream forming a diagonal from 
square F5 in the northwest to square Beta i in the southeast. Consequently the 
squares which were possible to excavate were Beta i, Alpha i, Ai, A2, Bi, B2, B3, 




Fig. 2. The plan of excavation at Bellan Bandi Palassa showing the relative distances 
of the skeletal remains from the ground surface and from the limestone bedrock. 



Cr, C2, C3, C4, Di, D2, D3, D4, D5, El, E2, E3, E4, and E5. Digging was continued 
in each square until the limestone bed rock was encountered. This base was humped 
with an apex at the C-line and declinations at A to the south and E to the north of 
the site. The loci of the specimens in this midden are represented in Text-fig. 2. 

In some of the burials the corpse was flexed and lying upon its left side (specimens 
BP2/17, BP4/8, BP2/21) or right side (BP2/25, BP3/27-34). Other skeletons 
were flexed but supine (BP 3/i5a, BP2/i5b). Fractional burial and bag burial 
cannot be excluded from consideration, although difficult to establish. The par- 
ticular artifacts found in direct contact with the skeletons are as follows : with 
BP2/17 were three unpitted hammer pebbles, of which one was discoidal, plus a 
bone dart. A cluster of twelve small pebbles in close proximity to one another 
suggest that they had once been encased in a bag and placed at the head of the corpse. 
In cleaning this specimen from its soil matrix, the mandible of a Macaca sinica, 



HUMAN SKELETAL MATERIAL FROM CEYLON 141 

probably female, was discovered (Personal communication with Dr. W. C. Osman Hill 
in January 1961, London). With BP2/20 was uncovered the left molar of Melursus 
valaha. With BP2/21 were found unworked spherical pebbles some 60 mm. in 
diameter which perhaps were bola stones. The mandible of Hystrix leucurus leucurus 
lay across the right zygoma of the specimen. Specimen BP3/27-34 was associated 
with an unpitted pebble above the level of the skull and a block of quartz under the 
right humerus. 

During the excavation of 1956 the remains of some nine individuals were uncovered. 
In the decade prior to this discovery, Deraniyagala had found fragmentary skeletal 
remains of single specimens from the middens of neighbouring sites. The significance 
of these sites has been discussed by Deraniyagala (19566, 1956c, 1956^, 1957a, 
19576 : 8, 20, 1958a, 1958& : 64-71, 1959, 1960^, 1960C, 1962, 1963a, 19636), Clark 
(1961 : 189-190), Cole (1963 : 87) and Coon (1962 : 424-425). However, no thorough 
laboratory examination of the bones was undertaken until i960 and 1961 when the 
specimens were sent on loan to the British Museum (Natural History) in London. 
It was advantageous to conduct the programme of research at this institution where 
the present investigator had available comparative osteological material from several 
collections of Vedda specimens and where excellent opportunities for subjecting the 
specimens to various biochemical tests existed. 

The Nature and Condition of the Specimens 

An abbreviated list of the skeletal specimens described in this report is given in 
Table i. These are from the collection made during the 1956 season of excavation. 
There are, however, two fragments included in the collection sent to the British 
Museum (Natural History) which cannot be assigned to any of the numbered and 
catalogued specimens. These are : 

1. A fragment of right ischium labelled BP2/i7g found in Square B2 with speci- 
men BP2/17. 

2. A fragment of right scapula which includes the glenoid cavity and a portion 
of the axillary border. This fragment is unlabelled but is associated with BP3 127-34 
in Square D4. 

Two specimens were found at Bellan Bandi Palassa which have not been described 
by the writer but which are noted in Deraniyagala's report of the site. These are : 

1. Specimen BP3/11 from Square C2 which lay at a depth of 2 feet 6 inches below 
the surface and 2 feet above the limestone. Only the maxilla and femur are des- 
cribed (Deraniyagala 1958^:230, 233, 236-237, table i). 

2. Specimen BP3/i5a was excavated from Square Ci at a depth of 3 feet 8 inches 
above the limestone. A second skeleton, specimen BP3/i5b, was discovered 
beneath it and it has been described by Deraniyagala (1958a : 230, 233, 260, pis. 
11-13, table I, 1963^:92-97). The skull of BP3/i5b (Deraniyagala 1958a : 233, 
pis. II, 12) was sent to the American Museum of Natural History in New York where 
its restoration is now under way. This is the specimen called T-23-B by Coon 
(1962 : 424-425). Through the kindness of Dr. Harry L. Shapiro the writer was 
permitted to examine this specimen. 



142 



HUMAN SKELETAL MATERIAL FROM CEYLON 






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144 HUMAN SKELETAL MATERIAL FROM CEYLON 

Of the specimens examined by the present writer, two are deserving of special 
comment. These are : 

1. The mandible of BPa/iyi which was found some lo inches away from the 
calvarium of specimen BP2/17. This was regarded at the time of discovery as 
belonging to specimen BP2/17, but upon cleaning the calvarium from its matrix it 
became obvious that the wear patterns of the maxillary and mandibular dentitions 
were strikingly different for the two specimens. 

2. Deraniyagala (1958a : 232) mentions the presence of another skeleton in the 
vicinity of the deposit where specimen BP2/21 was found. Its skull lay in Square C4 
with BP2/21 and its post-cranial parts extended into Square C3. Elsewhere in his 
report Deraniyagala (1958a : 230, 233) notes the presence of a specimen uncovered in 
Square C3 at approximately the same level as BP2/21. This specimen in C3 is 
called BP2/28 and it is without a cranium. The question arises as to whether these 
post-cranial bones in C3 may not belong to specimen BP2/21 rather than to another 
individual. The bones of BP2/28, namely a left humerus, left talus, calcaneum and 
femur were not available for study by the present writer. Furthermore, the great 
difference in the appearance of the cranial fragments of specimen BP2/21 is such that 
they could be certified as belonging to a single individual only after they had been 
cleaned of their matrix and concretion and examined for the purpose of reconstruction. 
This question of the presence of one or two individuals in Squares C3 and C4 is 
further complicated by the fact that Deraniyagala (1958a : 232, 260, pi. 9, tables 1-2) 
notes the presence of certain post-cranial bones belonging to specimen BP2/21 — a 
scapula, humerus, tibia, femur, talus, calcaneum — which again were not observed 
by the writer. 

As the bones were encountered in progress of excavation they were coated with 
shellac and labelled and packed in cotton. Fragile pieces were plaster- jacketed in a 
position exactly as they were found in situ. The plaster jacket for the calvarium of 
BP2/17 was later removed from one side and shellac dissolved in alcohol was poiired 
into the mass of matrix and bone. Specimens that were placed into the plaster 
jackets were BP2/17 (calvarium and pelvis), BP2/21 (cranium), BP2/25 (cranium), 
BP3/i5a (skeleton). Photographs were taken of the specimens in situ and some 
measurements were made upon the bones before their complete excavation. Upon 
the return of the archaeologists to the laboratory at Colombo these specimens were 
superficially cleaned. No reconstruction was attempted, save for the mending of 
bones which had been broken as a result of their transport from the field. Laboratory 
measurements carried out at the Colombo Museum were of a limited nature since the 
specimens were not entirely removed from their plaster jackets. 

The reconstruction and anthropometric analysis of the specimens was undertaken 
by the present writer at the British Museum (Natural History) . All of the specimens 
required some restoration, but warping and erosional damage, rather than actual 
breakage of bone, have most severely altered their pristine condition. Because of the 
poor state of preservation of the series, a morphological description is of necessity 
more significant than the tabulation of the metrical data. Nevertheless, where a 
metrical analysis could be undertaken, a record was made of the quantitative values. 



HUMAN SKELETAL MATERIAL FROM CEYLON 145 

The anthropometry is based upon the particular reconstruction that was considered 
by the writer to be of greatest accuracy for each specimen. The reconstructions are 
intentionally impermanent in order that other scholars may realize their own 
interpretations of the data. The reconstructing media are acetone cement and 
plaster of Paris, both of which can be flaked or washed from the specimens with ease. 

The osseous remains were not found in a uniform state of mineralization, but all 
were hard, due to the chemical nature of the soil in which they were embedded : 
the water acidulated by the humus had dissolved the limestone over which it flowed 
and formed a protecting environment for the calcium content of the bones. The 
bones of the lowest level were almost resting upon the limestone outcrop. The 
effluvia of the corpses had caused the limestone to disintegrate into yellow flakes to 
a depth of 5 cm., and these flakes had been subsequently impacted into a solid mass 
partially adhering to the bones. 

In addition to the osseous remains this series contains 120 permanent teeth of 
which 52 belong to male specimens and 68 to female specimens. 

II DESCRIPTION 
The Methodology of the Metrical Analysis 

The method of measurement for each of the values listed in Table 2 is to be found 
in Martin & Sailer (1957 : 453-499, 520-595). In this compendium the standard 
procedures for each measurement are described and listed by number. That number 
relevant to each measurement in Table 2 is given in parentheses immediately after 
its name. Indices are based upon these measurements. Variations and additions 
to these procedures are the following : 

1. Auricular- Vertex Height is taken according to the method devised by Ranke 
with the use of the Mollison craniophor, as described by Hooton (1946 : 738). 

2. The Frontal Arc is the first segment of the Glabella-Opisthion Arc and differs 
from the Nasion-Opisthion Arc described by Martin & Sailer (1957, Measurement 25) 
in its selection of the initial landmark. In the absence of a well defined nasion for 
any skull in the Bellan Bandi Palassa series, the most anterior point of the glabella 
in the median-sagittal plane was substituted for the conventional landmark in this 
arc measurement. 

3. The External Palatal Arc is measured from alveolon on one side of the upper 
jaw to alveolon on the opposite side, the tape passing across prosthion and parallel 
to the anterior aspect of the dental arcade. 

4. Height of the Zygomatic Bone is the distance measured with the sliding 
caUper from the anterior superior border of the frontal process of the malar to its 
anterior inferior border at the point of articulation with the maxilla. 

5. Breadth of the Zygomatic Bone is the distance measured with the sliding 
caliper from the superior border of the maxillary process to the anterior inferior 
border of the frontal process. 

6. Breadth of the Frontal Process of the Zygomatic Bone is taken as its greatest 
anterior-posterior diameter. The sliding caliper is placed at the lev^el of the zygo- 
matico-facial foramen when this measurement is taken. 



146 HUMAN SKELETAL MATERIAL FROM CEYLON 

7. The Mental Foramen Diameter is taken with the shding cahper, the points 
being placed upon the medial-lateral borders and the inferior-superior borders of 
the foramen for the determination of its maximum size. 

8. Sternal Head Diameter is the maximum anterior-posterior dimension of the 
medial extremity of the clavicle taken with the sliding caliper. 

9. Corocoid Process Breadth is the maximum anterior-posterior diameter of the 
corocoid process of the scapula, taken perpendicularly to the plane that the sliding 
caliper assumes in the measurement of Corocoid Process Length (Martin & Sailer 
1957, Measurement 11). 

10. Length of the Iliac Lines is measured with the sliding caliper from the anterior 
origin of the arcuate line on the pubic bone to its posterior termination at the sacro- 
iliac joint just superior to the pre-auricular sulcus. Both ilia are measured for 
this feature. 

In the discussion of the dentitions, the following abbreviations have been used : 
R = right side ; L = left side ; M3 = third molar ; M2 — - second molar ; Mi = 
first molar ; PM2 = second premolar ; PMi = first premolar ; C = canine ; 
I2 = lateral incisor ; Ii = central incisor ; the placement of the number above or 
below the line indicates that reference is made to either the upper or lower dentition, 
viz. RM3 = maxillary right third molar. 

Sex and Age Determination 

Specimen BP2/17 : This is a male who died between the ages of 25 and 30 years. 
The calvaria is the largest and most rugged in the series. The brow ridges are 
prominent and posterior to them is the trace of a frontal sulcus. The mastoid 
process is large with a supramastoid crest that exhibits a rough inferior margin. 
This robusticity of the mastoid is reflected also in the area of insertion for the splenius 
capitis and longissimus capitis. The digastric fossa is deep. The lateral border 
of the superior nuchal crest forms a moderately well defined ridge, but as its median 
portion is not represented in the fragment, its overall dimensions cannot be observed. 
Temporal lines are sharp at their frontal aspect and sweep posteriorly as an arc, of 
medium prominence on the parietals. The supramastoid crest is large. The 
zygomatic processes of the temporal are rough along their inferior margins, par- 
ticularly in the region of the anterior tubercle where some fibres of the masseter have 
their origin. The region of the attachments of zygomaticus major and minor on 
the malar shows prominent bossing. Medial to this is an elongated crest for the 
levator labii superioris and levator labii superioris alaeque nasi. The malar is short 
but massive. The maxilla exhibits very pronounced subnasal grooves. The 
alveolar arch of the palate is high. A basal view of the calvaria reveals further 
evidence of male robusticity : a deep mandibular fossa and a stout occipital condyle. 

The right and left innominates are characterized by narrow and deep ischiatic 
notches, large acetabulum, prominent sacro-iliac articulation, and a high upright 
ilium. A pre-auricular sulcus is present, but its dimensions are small and narrow. 
There is pronounced ridging of the region above the posterior superior iliac spine 
where gluteus maximus originates. The crest of the ilium is sharp, especially along 



HUMAN SKELETAL MATERIAL FROM CEYLON 147 

the line of origin of obliquus abdominis externus. The iUum is thick and heavy 
and the pelvic basin is small. Male characteristics are exemplified in the shoulder 
girdle with its relatively large clavicle and large scapular glenoid fossa and in the 
lower extremities where the linea aspera of the femur forms a pronounced pilaster. 

The absence of the pubic bones precludes the possibility of aging this specimen on 
the basis of progressive changes of the symphysial region. The dentition of the 
maxilla shows that the third molars have erupted and, like the other teeth, have 
undergone only a slight degree of attrition. Suture closure as an age indicator is of 
dubious value for this specimen due to trauma it has received from erosional forces 
and crushing. However, a general impression of its pristine condition can be obtained 
in a limited number of instances. Of the coronal suture, pars bregmatica is still 
undergoing closure but pars pterica is advanced. Of the sagittal suture, partes 
verticis and lambdica are advanced. The bregmatic and obelionic parts are eroded. 
Pars lambdica of the occipital suture has begun closure, but pars asterica is irregular, 
being most patent at its inferior portion. The masto-occipital suture is half closed. 
The squamous portion of the temporal is patent, but the spheno-temporal is advanced. 
Both spheno-paiietal and spheno-frontal are commencing closure. Radiography of 
the vault confirms visual observation that arachnoid granulations are not present. 
These observations suggest an age at time of death for the specimen of between 
25 and 30 years. 

Specimen BP2/i7i : This is an adult male. Its mandible, in comparison with 
those associated with specimens sexed as females, is of greater size, weight and 
thickness. The symphysis is higher, and the rami form a less obtuse angle in relation 
to the corpus. The short broad rami have robust pterygoid attachments at their 
gonia, which are thick and strongly everted. The mylo-hyoid ridge is pronounced. 

All of the teeth have erupted, and the third molars are moderately worn. The 
other molars reveal pronounced attrition. 

Specimen BP2/20-41 : This specimen may be that of an adult male, but criteria 
for sexing and aging are less certain than for other specimens of the series. Subnasal 
grooves are moderately developed. Of more masculine appearance is the mandible 
which is large and heavy with a pronounced mylo-hyoid ridge, very prominent 
pterygoid attachments, medium-sized genial tubercles and large digastric fossae. 

The dentition has erupted completely, and there is a moderate degree of abrasion, 
except for the upper left distomolar which is unworn. 

Specimen BP4/8 : This is an adult male characterized by robust musculature of 
the face and long bones. The malars are large and heavy, as is also the zygomatic 
process of the temporal. Subnasal grooves are pronounced, particularly in the 
incisive region. The palate is of moderate height with a large lump-shaped torus. 
The suborbital fossa is deep. Orbital and nasal borders are dull. 

The humeral supracondylar ridges are sharp, but the head of the humerus is 
reported by Deraniyagala as being comparatively small. The muscularity of the 
radius is pronounced as represented in the sharpness of the interosseous crest and 
the inferior border of the pronator quadratus attachment. The oblique line of the 
flexor digitorum sublimus muscle is well developed. Radial tuberosities are large. 



148 HUMAN SKELETAL MATERIAL FROM CEYLON 

The interosseous border of the ulna is less pronounced, but the groove where the 
extensor pollicis longus originates is marked. Likewise apparent is the insertion 
for brachialis on the anterior surface of the coronoid process. The supinator crest 
is high. The linia aspera of the femur forms a very prominent pilaster, and other 
well-developed areas of this bone are the crista hypotrochanterica and the trochanter 
minor. The femur is heavy and massive. The tibia is characterized by a moderate 
degree of muscularity. Its interosseous borders are sharp. Similarly the fibula 
has a sharp anterior margin, and the bone is deeply fluted. 

The teeth of this individual have completely erupted. The third molar has under- 
gone slight attrition, but the other teeth reveal a moderate degree of wear. 

Specimen BPi/6 : This is a female whose mandible exhibits signs of senile modifi- 
cation. The maxilla is delicate and the sub-nasal grooves are of medium develop- 
ment. The mandibular corpus is fragile and constricted at the molar region, but 
well developed, heavy and wide at the symphysial region. The chin form is median, 
and projection is pronounced. The mylo-hyoid ridge is of medium muscularity. 
The ramus is not robust, and its gonion is thin with reduced pterygoid attachments. 
E version of the gonia is medium. 

The teeth are erupted and, save for the slightly worn third molars, the remainder 
of the dentition shows a moderate to pronounced degree of wear. 

Specimen BP2/21 : This is a female whose age at time of death was between 18 and 
20 years. The supraciliary arches are reduced in development, and the superior 
orbital border reveals a delicate lipping. The mastoid process is medium in size, 
but the bossing for the sternomastoid is prominent. The mastoid shows a slight 
lateral projection. The occipital crests observed from their external aspects are 
reduced to traces. The supramastoid crest is moderately developed, but the temporal 
lines are traceable only on the frontal bone, where their conformation is rounded. 
The zygomatic processes and malars are small and gracile. For the maxilla the 
subnasal grooves are of medium development and the nasal sills are sharp. The 
palate is medium in height. Viewed basally, the calvarium presents a shallow 
mandibular fossa, small postglenoid process, and a minute stylo-mastoid foramen. 

The mandible is of medium size. The mylo-hyoid ridge is low, and the fossa 
for the mandibular gland is prominent. The gonia converge, but are strikingly 
thick and knobby as in specimen BP2/i7i, a male. The mandibular condyles are 
small and converging, and the coronoid process is high. 

The degree of muscularity of the post-cranial bones is within the moderate category. 
The humerus has a well-marked bicipital groove and tuberosity. The clavicle is 
sharply ridged in the region of the deltoid attachment. It is the low muscularity 
of the calvaria, rather than the features of the mandible and post-cranial bones, that 
suggests that the specimen is a female. 

The criteria for age determination are more satisfactory for this specimen. The 
third molars of the maxiUa and mandible have not completed their eruption and he 
partially embedded in their alveoli. Save for the lower incisors, attrition is neghgible 
for all of the teeth. Further proof of the young adulthood of this specimen is 
adduced by the metacarpal bones of the right hand where the epiphyses of the heads 



HUMAN SKELETAL MATERIAL FROM CEYLON 149 

and basal condyles have not completed ossification with the shaft. This fusion 
normally terminates in the twentieth year of life. Finally, sutural closure of the 
calvaria gives confirming data. The partes bregmatica and pterica of the coronal 
are commencing closure, but pars complicata remains patent. Of the sagittal 
suture, pars bregmatica is patent while the partes verticis, obelica and lambdica 
are commencing closure. The lambdoid suture is open at all regions save for pars 
media where closirre has started. The masto-occipital suture is patent as are the 
spheno-parietal, spheno-frontal, and spheno-temporal sutures. The squamous 
portion of the temporal, however, is closed, which is an artifact of the degree of 
preservation of the specimen. 

Specimen BP2/25 : This is a sub-adult female under 18 years of age. The sex 
criteria are less certain for this specimen than for the others of the series, but the 
cranium is small and its muscularit}^ is reduced. Unfortunately the frontal and 
occipital tori can not be observed. There is a moderate degree of ridging of the 
squamous portion of the temporal. The supramastoid crest is of moderate develop- 
ment, but the zygomatic process of the temporal is very thick. Nasal sills are sharp. 
Subnasal grooves are pronounced. The mandible is small and has a short corpus 
with little muscular development. The ramus is short and narrow with negative 
gonial eversion. 

Muscularity is very much reduced on the clavicle which is smooth and gracile. 
The radius is moderately fluted, but the interosseous crest is high. The radial 
tuberosity is low. The pilaster of the femur is of medium development and is 
mound-.shaped. The fossa hypo-trochanterica is deep. For the tibia, the anterior 
ridge is sharply defined but overall muscularity is reduced. 

Age is established on the basis of the unerupted state of the third molars of maxilla 
and mandible. Due to the inferior condition of the specimen the degree of suture 
closure on the cranium cannot be determined. Epiphyses of the long bones are 
complete, and this specimen can most accurately be aged as a sub-adult. 

Specimen BP3/27-34 : This is a female whose age at time of death was between 
30 and 35 years. 

The cranium is of medium size and exhibits a low degree of muscularity. The 
frontal torus is hardly discernible and glabella is very low. The occipital crests 
are reduced to low mounds. Inion is absent. Temporal crests are sharply defined 
on the frontal bone, but disappear in their progress posteriorly over the parietals. 
The anterior and posterior portions of the supramastoid crest are smooth. The 
malars are small and smooth. The temporal fossa has a moderate degree of roughen- 
ing, particularly on the posterior sphenoidal surface. The alveolar region is small, 
but subnasal grooves are pronounced and the palate is high and moderately ridged. 
The palatine torus is mound-shaped. Orbital and nasal borders are sharp. The 
suborbital fossa is deep and massive. The basal aspect reveals a deep mandibular 
fossa and a thick postglenoid process of a length unusual in females. The petrous 
portion of the temporal is small. 

The mandible is medium in size and light in weight. The corpus exhibits a 
pronounced mental spine emerging from a sharply pointed and projecting median 



I50 HUMAN SKELETAL MATERIAL FROM CEYLON 

protuberance. Genial tubercles are well defined and of medium size, but digastric 
fossae are small. The mylo-hyoid ridge is low. The angle formed by the ramus is 
more obtuse than that for the other mandibles of the series. The ramus is moderately 
broad and the gonia, while thin, are markedly crested for the attachment of the 
pterygoid and exhibit pronounced eversion. The coronoid process is high; the 
condylar neck is short. 

The small clavicle is thin and delicate. The scapular is small, and muscular 
attachments are weakly developed. The humerus is likewise reduced in robusticity, 
although the bicipital groove is well defined and has a sharp lateral lip. The small 
and sinuous radius has a large ulnar notch and an extensive but low radial tuberosity. 
The volar aspect of the radius inferior to the anterior oblique line shows a deep 
hollow for the reception of the belly of flexor pollicis longus. The styloid is of 
medium size. The ulna has a more prominent styloid process, and its crests are 
sharper. 

An estimate of age based upon suture closure alone would place this specimen 
within the range of 30 to 40 years at time of death. Conditions described below as 
patent or advanced are unreliable, as criteria of age in this specimen, due to absorption 
of the majority of the sutural margins, the post-mortem separation of the sutures 
as a residt of dessication and the consequences of the pressure of the over-burden 
upon sutural areas apparently complete in their closure. Hence partes bregmatica, 
complicata and pterica of the coronal are patent due to erosive factors working on 
the bone. For the sagittal, pars verticus appears to have been advanced in its 
closure but the degrees of fusion for pars lambdica, obelica and bregmatica are 
uncertain due to post-mortem separation of the parietals. The situation is the same 
for the lambdoid suture where the margins are preserved but unarticulated. Those 
sutures which have maintained their pristine condition are the spheno-parietal 
which is completely closed and the spheno-frontal which is advanced. The squamous 
portion of the temporal is fused to the sphenoid, but its relationship to the parietal 
cannot be accurately observed, save for a view of its superior portion on the right 
aspect of the vault where closure is well advanced. 

The dentition shows pronounced attrition. The upper third molars have erupted, 
but the status of the lower third molars is uncertain, for these teeth are not present. 

The Metrical and Morphological A nalysis of the Osseous Remains 
The Cranial Skeleton. The Calvarium (Table 2). All four calvaria of the 
series are dolichocranic, the female specimens being the narrower. There is less 
agreement in the indices of the Auricular Height of the vault in relation to Cranial 
Length and Cranial Breadth : the male specimen is hypsicranic and acrocranic, the 
female specimen is chamaecranic and tapeinocranic. However, if the Basion-Bregma 
Height is employed as a component of these cranial indices, the values for the male 
specimen fall within the same categories as do those of the female. Such a dis- 
crepancy in the indices of Cranial Height is due to the imperfect preservation of the 
basion region of specimen BP2/17. Therefore, the indices which utilize the auricular 
values are those which are preferable as the more reliable. 



HUMAN SKELETAL MATERIAL FROM CEYLON 151 

Cranial capacity cannot be directly measured for the specimens, and four formulae 
have been employed for the estimation of endocranial size. The calculation of 
von Bonin, while originally devised for male crania of natives from New Britain, 
has been demonstrated as suitable for male Australian crania as well (Hambly 
1947 : 35-39). Isserlis (1914) worked out a formula for estimating the capacity 
of crania from the Gaboon area of West Africa, and Hambly has noted its applicability 
to male and female crania of Vedda and Australian populations (Hambly 1947 : 57). 
However, when Isserlis' formula is applied to specimens BP2/17 and BP3/27-34 
of the Balangoda series, the results are 1589-72 cc. and 919-66 cc. respectively. 
Although there are known to exist two female Vedda crania with directly measured 
cranial capacities of 960 cc. (Hill 1941 : 90, 93-94), the value derived from Isserlis' 
formula in the cases under consideration is questionable. The formula of Lee & 
Pearson (1901) using Auricular Height confirms a low cranial capacity for specimen 
BP3/27-34. Without a confirmation of values for specimens of the Balangoda 
series that can be derived from direct measurement of cranial capacity, one is cautious 
in favouring the formula best suited to the series. These estimates do serve to 
illustrate the degree of variation between the large-headed male BP2/17 and the 
small, nannocranic female BP3/27-34. Differences in the values for the Cranial 
Module confirm this degree of variation. Specimen BP2/25, which was not recon- 
structed, appears to have had a small cranial vault and a greater delicacy of cranial 
architecture than BP3/27-34. 

The walls of the cranial vault are uniformly thick and heavy, due in part to their 
impregnation with mineral matter but to a greater degree this condition reflects 
their original nature. Porous areas are not observable. Their uniformly dense 
and grainy character is best observed radiographically. The frontal and anterior 
portions of the parietals are somewhat thinner than the bones of the posterior region 
of the vault. There is no sexual dichotomy in the thickness of the bones, although 
the parietals and occipitals of the female BP2/21 are the least massive in the series. 
Observed vertically, the conformation of the vault is sphenoid for BP2/17 and 
BP3/27-34 and rhomboid for BP2/21. In all three crania the greatest breadth is 
across the posterior third of the skull. The anterior portion of the vault of BP2/21 
differs from the others by the reduced proportions of its glabella and superorbital 
torus. Brow ridges are median in both female specimens. The male specimen 
exhibits a very large glabella and a heavy, divided frontal torus. The frontal 
development of BP3/27-34 falls between these extremes. Lateral aspects show 
a medium height of the frontal region but sexual differentiation in the frontal slope. 
The frontal bone of the male specimen inclines gradually from glabella to bregma 
with a curvature limited to the plane of the low frontal eminences. It has a large 
median boss. The females have a slightly bulbous frontal bone that curves evenly 
and smoothly to bregma, medium to pronounced frontal eminences, and no median 
crests. Specimen BP2/21 exhibits a median boss of modest size. Post-orbital 
constriction of the frontal is prominent in BP2/17 and BP3/27-34, but reduced in 
BP2/21. Frontal breadth is large for the male vault, small for the female. Supra- 
orbital foramina are large for both sexes. The frontal sinus is spacious in BP2/17. 

GEOL. II, 4 15 



152 



HUMAN SKELETAL MATERIAL FROM CEYLON 



The parietal region bears a sagittal crest of medium height in BP2/17 and BP2/21, 
but this region is not elevated in BP3/27-34. All the vaults of the series exhibit 
small post-coronal depressions. Parietal bosses are pronounced, and these lie well 
back towards the wider portion of the vault. From their apices, the sides of the 
brain case make a steep descent to the temporal fossae. Parietal foramina are not 
observable. The gradual curvature of the parietal portion of the vault, viewed 
from the lateral aspect, takes a sudden turn in a more vertical direction at lambda. 




Fig. 3. Left lateral contours of three Balangodese (BP2/17 

BP3/27-34 ). 



■, BP2/21- 




FiG 4. Frontal contours of three Balangodese (BP2/17 
BP3/27-34 ). 



- BP2/21 , 



HUMAN SKELETAL MATERIAL FROM CEYLON 153 

Lambda is moderately flat in all specimens but one, BP3/27-34. The occipital 
curvature is pronounced for all specimens. The occipital torus of BP2/17 is remark- 
able for its large size and robusticity : the female crania exhibit either reduction or 
no development of this region (Text-figs. 3, 4). 

The temporal region is full, particularly for the male specimen, and the sphenoid 
depression is remarkably large. The degree of cresting of the temporal lines is never 
pronounced along the parietals, but is marked on the frontals of the male vault. 
The squamous portion of the temporal is thick for BP2/17 and BP2/21, but thin for 
BP3/27-34. The size of the petrous portion is greater for the male. The auditory 
meatus is elliptical for the male, round for the females. The tympanic plate is 
consistently thick for both sexes. The mastoid is large for the male. Radiography 
reveals in the male specimen an extensive mastoid sinus. This consists of two 
polycameral sinuses in the superior aspect of the process along an anterior-posterior 
arc which traverses the width of the process. x\t its apex, a third polycameral sinus 
is visible. 

Sutures of the vault are simple in conformation for all specimens. Complexity 
is observed only in the peripheral margins of the coronal suture at pterion and of the 
sagittal at lambda. Wormian bones are not observable. Metopism is absent. The 
form of Pterion is H. 

Facial indices are ascertainable for specimen BP3/27-34. Its total facial index 
places it at the lower limit of the leptoprosopic class. The upper face is leptene. 
The nose is chamaerrhine. Whereas its orbits are chamaeconch, the orbits of 
BP2/17 are hypsiconch. The palates of all specimens are brachystaphyline. 

The angle of the upper face of BP3/27-34 places it within the orthognathous 
category as estimated by both the von Camper and Martin methods. The protrusion 
is of the alveolar sort rather than subnasal or total maxillary. Facial bones of the 
other specimens exhibit pronounced alveolar prognathism, save for the males 
BP2/17 and BP2/20-41 where alveolar protrusion is medium and slight respectively. 
The very pronounced alveolar prognathism of BP2/25 is to some degree an artifact 
of post-mortem distortion of the facial bones. Mid-facial prognathism is medium 
in the two cases where it is observable — specimens BP4/8 and BP2 127-34. 

The form of the orbit is rhomboid for the males and the female BP2/21, but square 
for the other female, BP3/27-34. Orbital inclination is 15° for BP2/17 and 16° 
for BP2/21, but the inclination is only 8° in the case of BP3/27-34. There is a 
striking sexual differentiation in the size and robusticity of the malars and zygomatic 
processes. Lateral malar projection is common to the females, but of the males, 
BP4/8 shows pronounced anterior malar projection. The total absence of nasal 
bones in the series precludes any comment as to the structure of this portion of the 
face save insofar as the maxillary walls and floor of the piriform aperture are preserved 
in BP4/8. From this evidence the nose appears to have been very broad with a 
large nasal spine and both sharp and dull sills. The nasal notch appears to have 
been deep for BP2/17 and BP3/27-34. Subnasal grooves of the incisive region are 
consistently prominent. The palate is elliptical except in specimens BP2/20-41 
and BP3/27-34 where the form is hyperparabolic, an appearance ascribable in part 



154 



HUMAN SKELETAL MATERIAL FROM CEYLON 



to the absence of third molars. The height of the palate is medium, but specimens 
BP2/17 and BP3/27-34 have deep palates. Palatine ridges are larger in males than 
in females and are mound-shaped rather than ridged for both sexes. The palates 
are unusual in their large overall size and in their considerable breadth (Pis. 1-7, 
10-14). 

Table 2 
Cranial Measurements and Indices 



Males 



Females 



Measurements 


BP2/17 


BP2/20-41 


BP4/8 


BP2/21 


BP2/25 


BP3/27-34 


Cranial Length (i) 


200 








183 




177 


Cranial Breadth (8) 


147 








133 




130 


Basion-Bregma Height (17) 


135 














Auricular- Vertex Height (21) 


132 












93 


Auricular- Bregma Height (20) 


127 












95 


Minimum Frontal Diameter {9) 


117 








no 




91 


Bizygomatic Diameter (45) 










121 




no 


Menton-Nasion Height (47) 










. , 




101 


Prosthion-Nasion Height (48) 










, . 


60 


64 


Nasal Height (55) 














44 


Nasal Breadth (54) 




23 










23 


Orbital Height-Right (52) 


37 








36 






Orbital Height-Left {52) 


37 












40 


Orbital Breadth-Right (51) 


41 














Orbital Breadth-Left (51) 


41 












30 


Biorbital Breadth (44) 


131 














Zygomatic Breadth-Right 


55 




52 


40 






Zygomatic Height-Right 


52 




44 


42 




47 


Zygomatic Frontal Process 


29 




22 


24 




22 


Breadth-Right 














Zygomatic Frontal Process 


25 












Breadth-Left 














Palate-External Length (60) 


61 






62 




53 


Palate-External Breadth (61) 


69 


70 




65 




63 


Palate-Internal Length (62) 


52 






50 




45 


Palate-Internal Breadth (63) 


36 


47 




42 




42 


Palate Height at M^ (64) 


19 


15 


14 


14 




18 


Arc-External Palate 


185 




152 








Arc-Transverse (24) 


355 








300 




270 


Arc-Glabella-Opisthion (25) 


355 








320 




285 


Arc-Frontal Length (25) 


132 








135 




115 


Arc-Sagittal Length (27) 


146 








152 




128 


Arc-Occipital Length (28) 


11 








53 




58 


Chord-Frontal Length {29-1) 


118 








120 




105 


Chord-Sagittal Length (30) 


130 








132 




112 


Maximum Circumference (23) 










520 




495 


Indices 














Cranial 


73-50 






72-68 


. . 


73-45 


Basion-Bregma Height-Length 


67-50 




. 


. 









HUMAN SKELETAL MATERIAL FROM CEYLON 



155 



Table 2 {continued) 
Cranial Measurements and Indices 

Males 



Females 





I 


^ f 


> 


Indices 


BP2/17 BP2/20-41 


BP4/8 BP2/21 BP2/25 


BP3/27-34 


Basion-Bregma Height- 


91-84 


. . 




Breadth 








Auricular-Vertex Height- 


66 • 00 


. . 


52-54 


Length 








Auricular- Vertex Height- 


89-69 


. . 


71-54 


Breadth 








Auricular-Bregma Height- 


63-50 


. . 


53-67 


Length 








Auricular-Bregma Height- 


86-53 


. . 


73-08 


Breadth 








Total Facial 


. . 


. . 


91-82 


Upper Facial 


. . 


. , 


58-18 


Nasal 




. . 


52-27 


Orbital-Right 


90-24 


. . 




Orbital-Left 


90-24 




75-00 


External Palate 


113-11 


104-84 


118-87 


Internal Palate 


69-23 


84-00 


93-33 


Cranial Module 


159-67 




133-33 


Cranial Capacity 








Basion-Bregma Height : von 


1448-75 


. . 




Bonin (1934 • 14) 








Lee & Pearson (1901 : 


1580-35 


. . 




225-264) 








Auricular Height : Lee & 


1775-83 


. . 


1098-87 


Pearson {Ibid.) 








Isserlis {1914 : 189) 


1589-72 


. . 


919-66 



The Mandible (Table 3). In contrast to the brachystaphaline upper jaw, the 
lower jaw is dolichostenomandibular. The alveolar portion of the mandible conforms 
to the characteristic shape of the palate, but the ramus describes an obtuse angle 
with the corpus, and moreover, is very broad. This results in a great Condylo- 
Symphysial Length but a moderate Bigonial Diameter. The corpus is thick and 
heavy at the symphysis, but is reduced in size, particularly among the females, as it 
sweeps posteriorly to the gonia. The chin is prominent and median for all mandibles, 
save for that of BP2/21 where the chin form is bilateral. The corpora of males are 
thicker and higher than the corpora of females, due in part to their more developed 
mylo-hyoid crests. There is less sexual diversity in the development of the digastric 
fossae and genial tubercles. These range in size from sub-medium to large. Alveolar 
prognathism is small in males and medium in females. The rami are robust in the 
males, their pterygoid attachments are well developed and gonial eversion is pro- 
nounced. Female rami are delicate and the gonia may be strikingly everted or 
convergent. The mandibular notch is small to medium, save for specimen BP2/21 
which has a deep notch due to the greater length of the coronoid process. In all 



156 



human skeletal material from ceylon 

Table 3 

Mandibular Measurements and Indices 



Males 



Females 



Measurements 

Condylo-Symphysial Length (68) 

Bicondylar Breadth (65) 

Symphysial Height (69) 

Bigonial Diameter (66) 

Height of Ascending Ramus — Right 

(70) 
Height of Ascending Ramus — Left (70) 
Minimum Breadth of Ascending Ramus 

—Right (71) 
Minimum Breadth of Ascending Ramus 

Left (71) 
Maximum Breadth of Ascending 

Ramus — Right (71-1) 
Maximum Breadth of Ascending 

Ramus — Left (71-1) 
Bimental Diameter (67) 
Depth at PM^— Right (69-2) 
Depth at PM^— Left (69-2) 
Depth at M'— Right (69-2) 
Depth at M^— Left (69-2) 
Depth at M^— Right (69-2) 
Depth at M^— Left (69-2) 
Depth at M^— Right (69-2) 
Depth at M^— Left (69-2) 
Thickness at Mi— Right (69-3) 
Thickness at M' — Left (69-3) 
Thickness at M^— Right (69-3) 
Thickness at M^— Left (69-3) 
Thickness at M^ — Right (69-3) 
Thickness at M^ — Left (69-3) 
Mental Foramen Diameter — Right 
Mental Foramen Diameter — Left 
Mean Angle (79) 

Indices 

Mandibular 

Fronto-Gonial 

Zygo-Gonial 









f 




^ 


BP2/i7i 


BP2/20 


-41 


BPi/6 


BP2/21 


BP3/27-3 


106 


79 






99 


93 


118 


"5 






105 


112 


28 


26 




31 


31 


28 


96 


no 






99 


lOI 




57 




70 


54 


49 


65 








48 


49 








30 


31 


31 


34 


35 






33 
36 


31 
41 


44 


•• 




•• 


36 


40 


45 


30 




. . 




41 


31 


27 








27 


31 


26 




27 




26 


28 


25 




27 


22 


23 


31 


26 




26 




23 


26 


25 




23 


23 


23 


28 


26 




25 


23 


20 


26 






21 


24 


20 


29 


27 




22 


24 


20 


16 


II 




14 


12 


ID 


16 


12 




15 


13 


9 


20 


12 




18 


15 


14 


19 


13 




17 


16 


13 


18 


12 




17 


17 


II 


19 


16 




18 


17 


II 




4x3- 


5 


15x2-5 








4x3- 


5 


25x3 






118 


123 






116 


133 


89-83 


68 


69 




94-28 
104-76 

81-82 


83-03 

81-25 

ioi-8i 



mandibles the condylar process does not rise very high above the coronoid process 
and the mandibular head is small. The lingula of the mandibular fossa is uniformly 
small. The ramus is broad and moderately high. The Balangodese mandible is 
of medium size but is distinctive for its broad and elevated rami and thick corpora. 
Sexual differences are difficult to isolate (Pis. 8, 9). 



HUMAN SKELETAL MATERIAL FROM CEYLON 157 

The Post-Cranial Skeleton. The Skeleton of the Trunk. Vertebrae were 
available for two specimens : BP2/17 and BP2/25. Their fragmentary and badly 
warped condition renders a metrical description of dubious value, but measurements 
were feasible for the fourth and fifth lumbar vertebrae of BP2/17. The cervical 
vertebrae are small and delicate with cordiform bodies and triangular neural foramina. 
The transverse processes are short. The atlas has a thin posterior arch and a very 
reduced posterior tubercle. The facet for the odontoid process is proportionately 
large for the accommodation of the thick, bulbous axial dens. The thoracic vertebrae 
are small. Their vertebral foramina are round and their bodies contain broad 
costal facets. The transverse processes are bulbous. The spinous processes are not 
large. In contrast to the pre-lumbar vertebrae, the last five segments of the spine 
are robust and are equipped with prominent posterior and transverse processes. 
The kiolorachic lumbar vertebrae of BP2/17 exhibit thick centra and wide epiphysial 
rings. For the fifth lumbar vertebra the Vertical Ventral (i) and Vertical Dorsal 
(2) Diameters are 23 mm. and 27 mm. respectively : the Depth of the Body is 30 mm. 
for the anterior-posterior diameter (4) and 46 mm. for the lateral diameter (7). 

Rib fragments accompany the vertebral remnants but none of the ribs is complete. 
The ribs of BP2/17 are very large at their sternal ends but never exceed 6-0 mm. in 
thickness. The costal grooves are deep with sharp borders. The ribs of the female 
specimen attain their greatest size at a point some distance lateral to the sternal 
head. The first rib shows a well-marked groove for the subclavian vessels, but the 
ribs just inferior to it are smooth and their thickness averages 3-0 mm. For both 
sexes the neck of the rib is robust and massive. 

The sternum of specimen BP2/17 is lacking its manubrium, but the body measures 
96 mm. in length and 32 mm. in breadth. Fusion of the sternal components is 
complete. Facets for six sternal articulations are well marked. Curvature is very 
slight. There are no sternal foramina. 

The Skeleton of the Upper Extremity (Table 4). The clavicle of the male specimen 
is large and shows a moderate degree of curvature. The sternal head is oval in form. 
The acromial head is moderately flat. Muscularity is reduced. In contrast to this 
condition are the clavicles of the female specimens which are small, delicately built, 
and with a medium to pronounced degree of curvature. Muscularity is much more 
apparent among the female clavicles which are characterized by sharp ridges for the 
deltoid attachment. These also show wide grooves for the subclavian vessels. Like 
the male clavicles, those of the females have oval sternal heads and flat acromial heads. 

The scapula is medium in size for the male, small for the female. For both sexes 
this portion of the upper extremity is robust. For the male specimen there is a 
pronounced obUquity in the direction of this spine in relation to a line tangent to 
the vertebral border. Since this border has undergone damage at the region inferior 
to the spine, the spino-vertebral angle can only be inferred and cannot be qualitatively 
or quantitatively ascertained. The axillary borders of both male and female 
scapulae are thick and tend to double their thickness in the vicinity of the glenoid 
fossa. The superior border is narrow and is lacking a scapular notch. The angle 



158 



HUMAN SKELETAL MATERIAL FROM CEYLON 



Table 4 
Measurements and Indices of the 
Bones of the Upper Extremity 
Males 



Females 
■«■ 



Clavicle 

Measurement 

Maximum Length — Left (i) 

Mid-Shaft Circumference — Right (6) 

Mid-shaft Circumference — Left (6) 

Mid-shaft Diameter — Anterior-Posterior 

—Right (5) 
Mid-shaft Diameter — Anterior-Posterior 

—Left (5) 
Sternal Head Diameter — Right 
Sternal Head Diameter — Left 
Index 

Length — Thickness — Left 
Scapula 
Measurement 

Morphological Length — Left (2) 
Corocoid Process Length — Left (11) 
Corocoid Process Breadth — Right 
Corocoid Process Breadth — Left 
Acromion Process Length — Right (10) 
Acromion Process Length — Left (10) 
Acromion Process Breadth — Right (9) 
Acromion Process Breadth — Left {9) 
Glenoid Fossa Height — Right (12) 
Glenoid Fossa Height — Left (12) 
Glenoid Fossa Breadth — Right (13) 
Glenoid Fossa Breadth — Left (13) 
Humerus 
Measurement 

Maximum Length — Left (i) 
Bicondylar Length — Left (2) 
Bicondylar Breadth — Right (4) 
Bicondylar Breadth — Left (4) 
Maximum Head Diameter — Right (9, 10) 
Maximum Head Diameter — Left (9, 10) 
Mid-Shaft Diameter — Anterior- Posterior 

—Right (6c) 
Mid-Shaft Diameter — Anterior-Posterior 

—Left (6c) 
Mid-Shaft Diameter Lateral — Right (6b) 
Mid-Shaft Diameter Lateral — Left (6b) 
Mid-Shaft Circumference — Right (7a) 
Mid-Shaft Circumference — Left (7a) 
Index 
Robusticity — Left 



BP2yi7 BP4/8 BP2/21 BP2/25 BP3/27-34 



39 



28 



18 



87 
43 
14 
14 
53 
52 
30 
26 

36 
38 

28 



53 
52 



18 
18 
59 
59 



40 



123 
32 
33 
10 

10 

19 



29-46 



30 
20 



302 
298 



35 
35 
12 

12 

17 
17 



48 
9 60 



HUMAN SKELETAL MATERIAL FROM CEYLON 

Table 4 (continued) 

Measurements and Indices of the 

Bones of the Upper Extremity 



159 



Males 



Females 



t 


% 


( 




> 


BP2/I7 BP4/8 


BP2/21 


BP2/25 


BP3/27 


Radius 










Measurement 










Maximum Length — Left (i) 


248 


. . 


. . 


. , 


Least Circumference — Left (3) 


38 




35 


38 


Maximum Head Diameter — Right (4-1, 


22 




•• 


•• 


5-I) 
Maximum Head Diameter — Left (4-1, 


22 


•• 


18 


19 


5-1 ) 

Mid-Shaft Diameter — Anterior-Posterior 


II 




15 


9 


—Left (5) 










Mid-Shaft Diameter — Lateral — Left (4) 


16 




19 


18 


Uhia 










Measurement 










Maximum Length — Left (i) 


261 






259 


Least Circumference — Left (3) 


33 


, , 


, . 


25 


Mid-Shaft Diameter — Anterior-Posterior 


15 


, , 


, , 


12 


—Left (11) 










Mid-Shaft Diameter — Lateral — Left (12) 


II 






8 


Capitate 










Measurement 










Length (i) 


. 


25 






Breadth (2) 


. . 


20 


, , 


, , 


Height (3) 


. 


18 






Trapezium 










Measurement 










Length (i) 


. 


23 






Breadth (2) 


. . 


19 


. , 


. . 


Height (3) 




II 






Scaphoid 










Measurement 










Length (i) 


. , 


23 






Breadth (2) 


. . 


13 


. , 


. . 


Height (3) 


. 


8 






Metacarpals 










Digit I Length (2) 


. 


71 






II 


. 


71 






„ HI „ 




68 






„ IV „ 


■ ■ . 


68 


, , 




„ V „ 


. 


48 






Phalanges 










Digit I „ (3) 


. . 


31 


. , 


, , 


,. IV „ 


. 


38 







i6o HUMAN SKELETAL MATERIAL FROM CEYLON 

of the glenoid fossa to the scapular body lies in a plane slightly lateral and superior 
to it. In form the fossa is elliptical, the inferior half being broad and the superior 
half narrowly constricted. Lipping of the fossa is submedium. The acromion is 
long and rectangular. The clavicular facet is unlipped. The corocoid process is 
fiat and extensive. 

The humeral shaft is long and only shghtly curved. A transverse section shows a 
prismatic pattern in the male specimen, a plano-convex pattern in the female. 
Muscularity is pronounced : the tuberosities are large, the bicipital groove is deep, 
the supracondylar ridges are sharp on medial and lateral aspects of the bone. The 
head is medium in size. The olecranon fossa is shallow and unperforated. Supra- 
condylar processes are absent. 

The radius is large for specimens BP4/8 and BP2/25, but small and sinuous for 
BP3/27. The shaft is straight in the male, but shows moderate anterior-posterior 
bowing in the female. The transverse section is oval in these bones for both sexes. 
The extremities of the radius are more massive for the male, but both sexes have 
shafts with well-marked muscular attachments. Tuberosities are extensive and 
low in females, prominent in males. The neck is longer for the male. Lipping of the 
capitulum occurs in the bones of both sexes. The styloid process is particularly well 
marked for males. Both sexes exhibit prominent muscular attachments on their radii. 

The ulna is triangular in transverse section. Curvature of the shaft is slight, 
occurring in the male only to a hmited degree in the region of the brachialis attach- 
ment just inferior to the coronoid process. Muscularity is pronounced for the male, 
moderate for the female. Both sexes show a prominent styloid process. The 
interosseous space is large as a result of radial rather than ulnar curvature. 

The bones of the hand tend to be large. The crests of the carpals are weakly 
developed. The form of the capitate is square. The scaphoid assumes a dumbbell 
conformity, but the trapezium is prominently ridged and metacarpals irregular. 
The metacarpals are long and exhibit pronounced dorso-ventral curvature. This 
being the hand of a sub-adult individual, the heads of the shaft are ununited. The 
metacarpal formula is II > III > IV > V(?) > I(?). The fifth metacarpal is not 
available for examination. 

The Skeleton of the Lower Extremity (Table 5). The femur of BP2/17 is stenomeric, 
and femora of the other male and the female of this series are eurymeric. Transverse 
section pattern is either round or quadrangular for the males but plano-convex for 
the female. The linea aspera is marked with a prominent pilaster, particularly in the 
case of the males. The shaft is uniformly thick and bowing is slight to medium in its 
degree. Tortion is slight to medium in its degree. Tortion is within the medium 
category (io°-20°), save for specimen BP2/17 where the tortion is more pronounced. 
The crista hypotrochanterica is of medium development for BP4/8, but the fossa 
hypotrochanterica is deep for all specimens. Trochanters major and minor are 
large and smooth. The neck of the femur is short and narrow and supports a small 
head. The intercondylar fossa is deep and broad in its transverse plane. There is a 
pronounced inferior orientation of the medial condyle and a slight anterior prominence 
of the lateral condyle. The adductor tubercle is reduced. 



HUMAN SKELETAL MATERIAL FROM CEYLON 



i6i 



Table 5 

Measurements and Indices of the 
Bones of the Lower Extremity 

Males 



Femur 

Measurement 

Maximum Length — Right (i) 
Bicondylar Length — -Right (2) 
Mid-Shaft Circumference — Right (8) 
Mid- Shaft Circumference — Left (8) 
Sub-Trochanteric Diameter — Anterior- 
Posterior — Right (10) 
Sub-Trochanteric Diameter — Anterior- 
Posterior — Left (10) 
Sub-Trochanteric Diameter — Lateral- 
Right (9) 
Sub-Trochanteric Diameter — -Lateral- 
Left (9) 
Mid-Shaft Diameter — Anterior- Posterior 

Right (6) 
Mid-Shaft Diameter — Anterior-Posterior 

—Left (6) 
Mid-Shaft Diameter — Lateral — Right (7) 
Mid-Shaft Diameter — Lateral — Left (7) 
Maximum Head Diameter — Right (18, 19) 

Indices 

Platymeria — Right 
Platymeria — Left 
Middle— Right 
Middle— Left 
Pilastric — Right 
Pilastric — Left 
Robusticity — Right 

Tibia 

Measttrement 

Mid-Shaft Diameter — Anterior-Posterior 
—Right (8) 

Mid-Shaft Diameter — Anterior-Posterior 
—Left (8) 

Mid-Shaft Diameter — Lateral — Right (9) 

Mid-Shaft Diameter — Lateral — Left (9) 

Nutrient Foramen Diameter — Anterior- 
Posterior — Right (8a) 

Nutrient Foramen Diameter — Anterior- 
Posterior — Left (8a) 

Nutrient Foramen Diameter — Lateral — ■ 
Right (ga) 



BP2/17 BP4/8 



26 



33 



Females 
BP2/25 



419 








•• 


414 










72 




80 

79 




66 

71 


25 




24 
26 




20 


20 




27 
26 




23 


24 




25 




24 


25 




27 




25 


20 




23 




18 


21 




23 




19 


35 










125- 


00 


88- 
100 ■ 


89 
00 


86-96 


83' 


•33 


92- 


GO 


75- 00 


84. 


•00 


85' 


•18 


76-00 


I20- 


•00 


108 ■ 


■59 


133-33 


iig- 


■05 


II7' 


•39 


131-59 


ID 


63 









26 


27 


23 


25 


27 


• 


19 


22 


16 


18 


21 


27 



19 



1 62 



HUMAN SKELETAL MATERIAL FROM CEYLON 

Table 5 [continued) 

Measurements and Indices of the 
Bones of the Lower Extremity 

Males 









<■ 

BP2 


/I7 


BP4/8 


Females 
BP2/25 


Nutrient Foramen Diameter — Lateral — 

Left (ga) 
Least Circumference — Right (lob) 
Least Circumference — Left (lob) 


19 

69 
66 




23 

72 


6r 


Index 














Middle— Right 
Middle— Left 
Platycnemia — Right 
Platycneraia — Left 






73- 

72- 

73- 


07 
00 

08 


81-48 

77-78 

69-60 


69-56 
65-52 


Fibula 














Measurement 














Mid-Shaft Diameter- 

— Left (3-2) 
Mid-Shaft Diameter- 
Least Circumference- 
Distal Breadth— Left 


— Anterior-Posterior 

-Lateral — Left (3-1) 
-Left (4a) 
(4-2) 






15 

9 
33 
23 


•• 


Talus 














Measurement 














Length— Right (i) 
Breadth— Right (2) 
Height— Right (3) 






45 
32 
18 




' * 


•• 


Calcaneum 














Measurement 














Length— Right (r) 
Breadth (2) 






67 
37 




:; 


:: 


Cuboid 














Measurement 














Length— Left (i) 
Breadth— Left (2) 






35 
29 




•• 


-• 



The tibiae are eurycnemic for specimen BP2/17 and mesocnemic for BP4/8 and 
BP2/25, although these latter two stand at the upper and central loci of the mesoc- 
nemic category. Tibial lengths could not be ascertained. The posterior half of 
the transverse section of the shaft is oval, but in BP4/8 the conformation is closer 
to plano-convex. The bones are heavy and thick for the males. Bowing of the 
shaft is very slight. Muscularity is moderate for both sexes. 

The fibula is represented by the single fragment belonging to BP4/8. It is the 
left fibula. The form of its transverse section is triangular. Whereas the anterior 



HUMAN SKELETAL MATERIAL FROM CEYLON 163 

border is sharp, fluting of the shaft is sub-medium in development. The lateral 
malleolus is medium in size and its articular facet is extensive. The shaft is straight. 
The talus possesses a large triangular articular facet for the fibular malleolus. 
The groove for the flexor hallicus longus is medium in development. The neck of 
the talus is not constricted. The calcaneum is short and deep with a round posterior 
curvature of the sustentaculum tali. The cuboid exhibits pronounced curvature of 
the cuboid-lateral cuneiform facet. The articular facet is large, round, and its 
borders poorly defined. There is no lipping of the facet. The cuboidal ridge is 
prominent. 

Table 6 

Measurements and Indices of the Bones of the Pelvis 

Male 
BP2/17 
Sacrum 

Measuyement 

Anterior Length (2) 102 

Sacral Breadth (5) 125 

Lateral Diameter of Body i (19) 46 

Anterior-Posterior Diameter of Body i (18) 29 

Anterior Height of Body i (24) 29 

Anterior Height of Body 2 (24) 28 

Index 

Sacral 122-55 
Innominate 
Measurement 

Innominate Length — Right (i) 207 

Innominate Length — Left (i) 184 

Innominate Breadth — Right (6a) 141 

Innominate Breadth — Left (6a) 142 

Distance from Acetabulum to Apex of Ilium — Right (9) 132 

Distance from Acetabulum to Apex of Ilium — Left (9) 114 

Length of Iliac Lines — Right 262 

Length of Iliac Lines — Left 260 

Vertical Diameter of Acetabulum — Right (22) 42 

Horizontal Diameter of Acetabulum — Right (22) 37 

Breadth of Sciatic Notch — Right (8) 50 

Distance from Anterior Superior to Posterior Superior 138 
Iliac Spines — Right (12) 

Sacrum and Innominates 

Measurement 

Bi-Uiac Diameter (2) 218 

Sagittal Diameter of the Pelvic Inlet (23) 157 

Transverse Diameter of the Pelvic Inlet (24) 107 

Index 

Pelvic Inlet 146-81 

Innominate Breadth-Height — Right 129-58 



i64 HUMAN SKELETAL MATERIAL FROM CEYLON 

The Skeleton of the Pelvis (Table 6) . The single sacrum observable in the series 
is platyhieric. The sacral type is homobasal. Of the five sacral vertebrae the first, 
second and third articulate with the innominates. Curvature commences at the 
second sacral body but is not pronounced. Spina fissa is absent. The overall size 
of the sacrum is large. Deraniyagala considers the Stone Age population of Ceylon 
to have been steatopygous. He discusses this in relation to the small pelvis of the 
female skeleton found at the site of Alu galge (TeluUa) (Deraniyagala 19556 : 301) 
and considers the fossils from Bellan Bandi Palassa to have exhibited the same trait 
(Deraniyagala 1958a ; 255). The present writer is unable to find any evidence 
either to confirm or reject this opinion. 

The innominates of this individual are thick and massive. Their size is moderate 
and the muscular attachments are prominently sculptured. The ilium has heavy 
and irregular crests along its superior rim and deep hollowing of the iliacus portion 
of the medial surface. The origin of gluteus maximus is well marked. There is 
considerable torsion between ilium and ischium. The anterior superior iliac spine is 
broad and blunt. The ischia are divergent in their orientation and are moderately 
elongated in size. The ischiatic notch presents a higher angle than is usual in male 
specimens. The pelvic inlet is cordiform and its index places it within the dolichopel- 
lic group, although the absence of the complete pubic area renders this statement 
subject to question. 

An Estimation of Stature (Table 7) . Limb proportions are unfortunately impossible 
to establish for the Balangoda population due to the paucity of long bones complete 
enough for their maximum lengths to be accurately measured, and to the absence 
of any bones of the upper and lower extremities belonging to the same individual. 
Those long bone lengths which could be measured are listed in the table below, 
where they are employed as components of the formulae devised by Trotter & 
Gleser (1952, 1958) for estimation of stature from long bones. 

The lengths of the bones of two males, BP2/17 and BP4/8, and one female, 
BP3/27-34, are employed in formulae appropriate to American Whites, American 
Negroes, and American Mongoloids, mainly Chinese. The stature estimates for 
each of the three groups are given. 

In an effort to ascertain which of the three formulae might be best suited to the 
Balangodese, the writer applied them to the long bone lengths of three Veddas 
whose stature in life was known and whose long bones had been measured by Hill. 
These individuals were two males — Burunda, specimen 1949-12-7-4, and Tissahamy, 
specimen 1949-12-7-6, both of British Museum of Natural History (Hill 1941 : 148- 
149, tables I, 22). The former individual was from Dambane and was partly 
Sinhalese ; the latter from Kalakoluebe near Yakkure was non-Sinhalese in racial 
background. The female Vedda available for comparison was not measured in Ufe 
but skeletal stature is known. This is the Vedda specimen in the osteological collec- 
tion of the Bombay Natural History Society (Hill 1941 : 227-235). 

Such estimations of stature as given here are only indicative of the order of stature 
which is to be anticipated for the three Balangodese individuals. For the Veddas, 



HUMAN SKELETAL MATERIAL FROM CEYLON 



165 



the formula for Mongoloids renders a value that most closely approximates the living 
stature of Burunda, but for Tissahamy, the formulae for the Negroes is a nearer 
approximation. 

The Metrical and Morphological A nalysis of the Dentition 

In their dental morphology the Balangodese have teeth which are moderate to 
large in size by general standards for post-Pleistocene hominid populations (Table 8). 
The order of size of the molars within the molar rows of each specimen show con- 
siderable variation. For BP2/17, the RLM^ and RLM^ are progressively smaller 
than the RLM^. This is also the case for the mandibular molar row of BPa/iyi, but 
in the remaining male specimens the maxillary molar row shows the first molars of 
greater size. The mandible of BP2/20-41, however, shows a larger third molar for 
the row. For the female BPi/6, the maxilla has a second molar of greatest size 
followed in descending order by the first molar and the third, but the mandibular 
molar row has the third molar larger than the first and the first larger than the second. 

Table 7 

An Estimation of Stature Based Upon 
Lengths of Long Bones for Balangodese and Veddas 



BALANGODESE 

Males 



BP2/17 



BP4/8 



Female 
BP3/27-34 



Length Stature 
Right 



Length Stature 
Left 



Length Stature 
Left 



White : 

Humerus 

Radius 

Ulna 

Femur 

Tibia 

Fibula 

Negro : 

Humerus 

Radius 

Ulna 

Femur 

Tibia 

Fibula 

Mongoloid 

Humerus 

Radius 

Ulna 

Femur 

Tibia 

Fibula 



248 
261 



419 



1627 



1734 
1713 



302 
259 



248 
261 



1663 



302 

259 



419 



1602 



1654 
1706 



1624 
1656 







302 


I64I 


248 


1698 






261 


1683 


259 


1676 



419 



1626 



i66 



HUMAN SKELETAL MATERIAL FROM CEYLON 



Table 7 (continued) 

An Estimation of Stature Based Upon 
Lengths of Long Bones for Balangodese and Veddas 

VEDDAS 









Mai 


^s 






Female 








A 








Bombay Nat. 




r 








^ 




1949- 


-12-7- 

A 


4 


1949 


-12-7-6 

A 




Hist. Soc. 




f 
Length 


Stature 


f 
Length 


Stature 


Length 


Stature 




R L 


R 


L 


Right 


R 


L 


R L 


\Vhite : 


















Humerus 


300 301 


1648 


1650 




. , 


250 


262-5 


1503 1540 


Radius 


241 239 


1707 


1700 






202 


202 


1560 1560 


Ulna 


266 263 


1732 


1721 






221 


221 


1566 1566 


Femur 


429 430 


1650 


1653 


398 


1579 


361 


362 


1493 1495 


Tibia 


365 368 


1703 


1710 


328 


1613 


3" 


311 


1572 1572 


Fibula 


358 355 
Mean = 


1686 
1686 


1678 


325 


1600 
1597 


301 


300 


1538 1535 


Negro : 


















Humerus 


300 301 


1619 


1622 






250 


262-5 


1475 1512 


Radius 


241 239 


1654 


1648 






202 


202 


1525 1525 


Ulna 


266 263 


1679 


1669 






221 


221 


1535 1535 


Femur 


429 430 


1823 


1825 


398 


1558 


361 


362 


1480 1482 


Tibia 


365 368 


1653 


1659 


328 


1572 


3" 


311 


1535 1535 


Fibula 


358 355 
Mean = 


1638 
1677 


1631 


325 


1561 
1564 


301 


300 


1505 1503 


Mongoloid : 


















Humerus 


300 301 


1636 


1638 






250 


262-5 


1502 1535 


Radius 


241 239 


1673 


1667 






202 


202 


1535 1535 


Ulna 


266 263 


1700 


1690 






221 


221 


1543 1543 


Femur 


429 430 


1648 


1650 


398 


1681 


361 


362 


1502 1504 


Tibia 


365 368 


1687 


1694 


328 


1598 


311 


311 


1558 1558 


Fibula 


358 355 
Mean = 


1665 
1667 


1658 


325 


1586 
1622 


301 


300 


1528 1526 


Known 


(Living) 1629 


(Livi 


ng) 1493 




(Skeletal) 1267 


Stature : 



















Specimen BP2/21 offers another variant : the maxilla shows a large first molar, a 
medium second molar and a smaller third molar, but in the mandible the largest 
tooth is the second molar followed in size by the first and then the third. Specimen 
BP2/27-34 has a larger maxillary second molar than the first molar, but a larger 
mandibular first molar. These data can be summarized as follows : 



Males : 



Females 



BP2/17 
BP2/i7i 
BP2/20-41 
BP4/8 

BPi/6 
BP2/21 



Maxilla. M3 > M2 > Mi 



Maxilla. 
Maxilla. 

Maxilla. 
Maxilla. 



Ml > M2 > M3 
Ml > M3 > M2 

M2 > Mi > M3 
Mi > M2 > M3 



BP3/27-34 Maxilla. Mi > M2 



M3 > M2 > Ml 
M3 > Mi > M2 

M3 > Mi > M2 
M2 > Mi > M3 



Mand. 
Mand. 

Mand. 
Mand. 
Mand. M2 > Mi 



HUMAN SKELETAL MATERIAL FROM CEYLON 167 

The molars are round in their conformation, but rectangular molars are present 
in the LM^ of BP2/20-41 and in all molars of BP2/21 save for its RLM^ which is 
round. Premolars are rectangular in the maxillae of both sexes, but variability 
occurs in the mandibles. Specimen BP2/20-41 has RLPMi and RLPM^ vi^hich 
are round ; another male, BP2/i7i, has a triangular LPMT although its other mandi- 
bular premolars are round. In both these cases, the molar pattern is round, with 
the exception of the LM^ of BP2/20-41 noted above. There is no maxillary dentition 
available for BP2/i7i. The female BP3/27-34 has a round RLPM^ adjacent to a 
rectangular RLPM2 and round molars. Canines of the maxillae and mandibles are 
triangular for both sexes, but rectangularity appears in the RLC of the male 
BP2/20-41. Incisors are uniformly rectangular. 

Cusp and groove patterns are not observable for all of the teeth of the series due 
to the high incidence of attrition. For the maxilla, the males show pronounced 
variability. Specimen BP2/17 shows 5 cusps for the LM^ and 3 cusps for RLM-. 
The other male, BP4/8, has 4 cusps on the RMi, RM^, and RM^, the other molars 
not being observable. In contrast, the maxillary molars of the female specimens are 
of the 4-cusp pattern. The RLM- of BP2/21 has 6 cusps. In the single male, 
specimen BP2/20-41, for which the cusps of the lower molars are observable, the 
LM3 has 5 cusps and the remaining left and right molars have 4 cusps. The three 
female specimens show one, BPi/6, with 5 cusps for the RLM^ and RLMT, but 4 
cusps for the RLM 2. Specimen BP2/21 has 5 cusps for all molars, save for the 
LM3 which has 6 cusps. For specimen BP3/27-34 only the RLM2 with 4 cusps is 
available. Of the four specimens for which both maxillae and mandibles are 
available, there is no case of correlation of cusp pattern except for specimen BP3/27- 
34 where the second molars are alone observable. The upper and lower premolars 
of females are bicuspid, the buccal cusp being the larger and higher. Male premolars 
are also of this pattern, save for the RLPM- of BP2/20-41 where the premolars have 
4 cusps due to the presence of accessory cusps on the distal inclination of the buccal 
cusp. 

Carabelli's cusp is present in the LRM^ of one specimen, BP2/21. The cusp is 
small and the mesio-lingual groove is of moderate development. 

Accessory cusps are found on the second and third maxillary molars of the male 
specimen BP2/17. They are small and occupy loci on the mesial half of the buccal 
surface. 

Crenulation or excessive wrinkling of the occlusal surface of the upper third molars 
is shared by males — BP2/17 and BP2/20-41 — and females — BPi/6 and BP2/21. 
It is absent in the remaining male maxillae and the third molars of the female 
BP3/27-34 are unavailable for examination. Its occurrence is marked by the 
presence of 5 cusps in the male maxillae. For the females, BPi/6 has the 4-cusp 
pattern and BP2/21 has 6 cusps. All cases are characterized by pronounced crenula- 
tion regardless of cusp number. The mandible of male BP2/20-41 shows a deeply 
crenulated LM^ with 5-cusp pattern — a reflection of the maxillary LM^^ of this same 
specimen. The mandible of female BPi/6 shows a pronounced crenulation of the 
RLM^ , but the cusps of these lower third molars constitute the 5-cusp pattern instead 

GEOL, 11,4 16 



i68 HUMAN SKELETAL MATERIAL FROM CEYLON 

of the 4-cusp pattern of the upper third molars of the same specimen. In specimen 
BP2/21 six cusps are observed in both maxillary and right mandibular third molars 
but the RM3 has 5 cusps. 

For the maxilla, cusp patterns are unusual in the dentition of female specimen 
BP2/21 where the RLMi and RLM^ are of the +3 type. The groove patterns for 
the remaining maxillary dentitions of the males are of the +4 pattern. The 
mandible of BP2/21 shows a RLM2 of the +5 pattern and a RLMi of the Y5 type. 
In specimen BPi/6 the RLMT is also of Y5 type, but the RLM2 is +4 and the RLMs 
is +5- The other female specimen, BP3/27-34, has a RLM^ of +4 type. The 
only male mandible which offers data on groove pattern is specimen BP2/20-41, 
whose LM3 is Y5 and whose RLM2 and RLMi are +4 in form. Taking the series 
collectively, it would appear that the basic cusp pattern for the upper molars is of 
the +4 type. The lower molars are characterized by the groove patterns Y5, +5, 
and +4. 

A correlation of cusp type and groove pattern reveals that the presence of 4 cusps 
is associated with the +4 pattern in the mandible as observed in specimens 
BP2/20-41, BPi/6, and BP3/27-34. However, the mandibular 5-cusp type may be 
associated with either the +5 type, as with the RLM^ of BPi/6 and RLM2 of 
BP2/21, or with the Y5 type as represented by the LM^ of BP2/20-41, RLMi of 
BPi/6, and RLMT of BP2/21. As is obvious from an examination of these cases 
for which a mandibular molar dentition is available, each specimen has its unique 
combination of cusp type and groove pattern. Nevertheless, where the M^ has 
5 cusps, its groove pattern is a Y5. The M^ is consistently a + pattern and in 
three of the specimens a +4, namely with BP2/20, BPi/6, and BP3/27-34. In one 
specimen, BP2/21, this is +5. Crenulation of the M^ is associated with the presence 
of 5 cusps which are either of Y5 pattern hke BP2/20 or a +5 pattern like BPi/6. 
The groove pattern of the M^ of BP2/21 is unobservable, but the cusp number is 
5 for the RM3 and 6 for the LM^, both molars showing crenulation. This point 
cannot be confirmed for the mandibular third molars of BP3 127-34. In the maxilla, 
as noted above, crenulation of the third molar is associated with 4 cusps (BPi/6), 
5 cusps (BP2/17, BP2/20-41) or 6 cusps (BP2/21). 

Attrition of the molars of male and female dentitions show considerable variation. 
The maxillary molars are very slightly worn in BP2/17. All cusps of the RLM^- 
and the mesio-labial cusp of the LM- exhibit abrasion of the enamel with consequent 
exposure of the dentin in the form of pits at the apices of the cusps. Direction of 
abrasion cannot be established for this specimen. In contrast to this condition 
are the heavily abraded molars of BP2/i7i. The RLM^ show the least wear for the 
six teeth of the molar rows, but like them a rim of enamel surrounds a lake of dentin 
from the centre of which protrudes a reduced cusp which is still covered with enamel 
in the RLM^, LM2, and LMi. Wear is greatest on the labial aspects of each molar, 
but the enamel rim is everjrwhere intact. The dentin surfaces are deeply hollowed 
around the central reduced cusps, the sculpturing being most pronounced on the 
buccal sides. The maxilla of BP2/20-41 contains a fourth molar on the left side 
which, hke the LM^ adjacent to it, shows an absence of attrition. The other 



HUMAN SKELETAL MATERIAL FROM CEYLON 169 

maxillary molars show a moderate degree of wear, the greatest degree of abrasion 
occurring on the buccal portions of the occlusal surface. The enamel rim is thin 
and brittle on the mesial portion of these molars, but thick and robust on the other 
portions. A similar degree of wear characterizes the mandibular molars, the LM^ 
showing less wear than the remaining molars, the buccal sides being the most abraded 
and denuded of enamel. The male BP4/8 shows slight wear of the RM^ and medium 
wear of the RM^ and RM^. The exposed dentin forms kidnej^-shaped lakes over 
the occlusal surfaces, and the enamel borders are stout. 

The RLM^ of female specimen BPi/6 shows very slight abrasion. The adjacent 
molars are moderately worn with small pits of exposed dentin on their cusps. The 
RLMi present kidney-shaped lakes of exposed dentin on the mesio-buccal occlusal 
surfaces. The direction of the abrasive force is lingual for both molar rows. The 
mandible of this specimen reflects the maxilla in regard to attrition, save that the 
RLM^ exhibits pronounced abrasion. The maxillary and mandibular molars of 
BP2/21 show negligible wear, save for the first molars which have developed small 
pits of exposed dentin on the apices of their cusps. Direction of this slight abrasion 
appears to be buccal. As the third molars are in process of eruption, their elaborately 
crenulated occlusal surfaces are untouched by attrition. The dentition of BP3/27-34 
is heavily abraded. The third molars have been lost ante-mortem, and the first and 
second molars exhibit occlusal surfaces denuded of enamel, save for an island of 
enamel forming a central cusp. Greatest wear occurs on the lingual surfaces. The 
enamel rims are narrow and brittle and enclose poorly sculptured lakes of the dis- 
coloured dentin. 

As with the molars, the premolars of females are more frequently abraded than 
are the premolars of males. For specimen BP2/17 the premolars are very slightly 
worn, although attrition of the buccal cusps has resulted in exposure of dentin on 
the RLPMi and RPM-^. The mandibular premolars of BP2/i7i are heavily abraded, 
the RPM2 exhibiting the greatest degree of attrition. The exposed dentin of these 
teeth lacks the central reduced cusps found on the molars. Orientation of the abra- 
sive force is buccal. The enamel borders of these premolars are rounded in contrast 
to the sharpened ridges that constitute the enamel borders of the molars. Maxillary 
and mandibular premolars of BP2/20-41 show a moderate degree of attrition. The 
force of abrasion appears not to favour either side of the maxillary or mandibular 
premolar row, but the distal portions have maintained the greater parts of the 
enamel, particularly for the RLPMi. The maxillary premolars of BP4/8 show 
medium attrition, the RPM-^ having the least amount of dentin visible. Wear is 
greatest for both RPM^ and RPM^ on the labial portion, although the lingual cusps 
are marked by several small pits and lines of exposed dentin. 

All maxillary premolars of female BPi/6 show a moderate degree of wear, except 
for LPMJ- which is heavily worn. Unlike the other premolars whose bays of exposed 
dentin are limited to buccal and lingual areas separated by mesio-distal ridges of 
enamel, the occlusal surface of LPMJ- is denuded of all enamel and its rim is sharp 
and narrow. This tooth is separated by a gap of 10 mm. from LPM^ which is due in 
part to post-mortem distortion of the maxilla, but which perhaps reflects as well an 



I70 HUMAN SKELETAL MATERIAL FROM CEYLON 

abnormal orientation of this tooth in the molar-premolar row. Unfortunately the 
mandibular premolars of BPi/6 are missing. Direction of wear appears to have had 
no preference for either side of the premolar rows. Specimen BP2/21 exhibits very 
slight attrition of both maxillary and mandibular premolars, although the RLPM2 
shows somewhat more abrasion than the others. Pits of dentin on the occlusal 
surface are numerous and are greatest in frequency on the labial portions of the 
premolars. In contrast are the maxillary and mandibular premolars of BP3/27-34, 
all of which are heavily worn, particularly on their labial sides where the enamel 
borders of the RLPM2 have broken down. Vestiges of the lingual cusps are visible 
on the RLPM^^. 

Canines show slight attrition in specimen BP2/17, the apices of the cusp being 
marked by very small pits of dentin. Those of BP2/20-41 are moderately worn 
with the greatest degree of attrition on the labial aspects. The RC of BP4/8 is 
characterized by a lingual and labial pit on its occlusal surface, but these are reduced 
in size. The maxillary LC of BPi/6 has moderate wear on its labial aspect. Those 
of BP2/21 are marked by numerous pits of exposed dentin but the cusps are not 
reduced in size. The canines of BP3/27-34 are very abraded in the maxilla and in 
the RC of the mandible, although the enamel borders are well preserved and rounded 
in form. 

Incisors are moderately to heavily abraded for all specimens. The RLI^ of BP2/17 
present a bar-like incisal margin of exposed dentin, portions of which are pitted. 
The more pronounced attrition of the LI-^- of BP2/20-41 shows a triangular mass of 
exposed dentin on the cutting surface, the enamel rim being reduced in size and very 
thin along its mesial aspect. The lingual side is worn to the base of the crown. The 
RI-^ and RI- of BPi/6 are moderately worn, the central incisor showing slightly 
greater wear. The exposed dentin is rectangular in form. The maxillary and 
mandibular incisors of BP2/21 are heavily abraded with their rectangular cutting 
surfaces. Due to post-mortem damage to the mandible, the RP and Ll2^ have 
become dislocated so that the RI2 has been pushed to the left side where it is adjacent 
to the LI 2. The cutting edges are rectangular. Pronounced wear characterizes 
the maxillary LI^ which is worn to a triangular-shaped stub of exposed dentin. 

Deraniyagala (1958a : 255, 260) remarks that the lower teeth are always more 
heavily abraded than the upper teeth. This is the tendency for the teeth taken as 
a whole, although for the few incisors available in the sample the upper teeth appear 
to be more heavily abraded. However, if the dentitions are examined according to 
specimen, only two, BPi/6 and BP2/21, illustrate Deraniyagala's claim. The other 
two specimens, BP2/20 and BP3/27-34, show an equal degree of wear for teeth in 
both jaws. There is a higher incidence of mandibular molar wear among males than 
females, but maxillary molar wear shows no sexual dichotomy. Females show a 
higher incidence of premolar attrition and maxillary canine attrition, but males have 
a higher incidence of wear among mandibular canines and maxillary incisors. No 
comparison by sex is possible for the lower incisors since these are lacking for male 
specimens. 

Shovel shaping of the incisors is lacking in the male dentitions available for 



HUMAN SKELETAL MATERIAL FROM CEYLON 171 

examination, but is present in the RI^ and RI^ of female BPi/6 and in the LI^ and 
LP and RLI^ and RH of the female BP2/21. It is expressed to a pronounced 
degree only in the maxillary incisors of BP2/21, the size of the shovel shaped depres- 
sions of this specimen's mandibular incisors being slightly developed. Development 
is medium in the BPi/6 incisors. The single case of a lingual tubercle occurs in the 
LI^ and RLI2 and RH of BP2/21. The tubercle is prominent in the mandible, 
but small in the maxilla of this female specimen although a foramen cecum incisal 
to the tubercle marks this feature with clarity. 

Some features that frequently appear in the dentitions of other Recent hominid 
populations are not observable in the Balangoda series. Caries and dental abcesses 
are not observed. Ante-mortem tooth loss is suggested in the case of a single 
specimen — the RLM^ of female BP3/27-34. The alveoli in this case appear completely 
absorbed. The maxillary third molars of this specimen appear to have been lost 
post-mortem, the RM^ showing a small alveolus with unabsorbed borders. Super- 
numerary teeth are found in the case of the LMi of the maxilla of BP2/20-41. 
The size of this distomolar tooth is reduced, and its form is of the compressional or 
rectangular type. It was never of any functional benefit. Buccomesial to the 
RM2 of specimen BP2/21 is a peg-shaped paramolar which is encased in enamel 
and is half as high as the adjacent teeth. Suppression is not a feature of the dentitions 
of the series. The upper and lower third molars of specimen BP2/21 were almost 
completely erupted at the time of death of this individual. 

Deraniyagala (i958<2 : 255) remarks that bite is of the edge-to-edge type. This 
cannot be ascertained with certainty for any of the specimens, although it may have 
been a trait for those individuals with heavily worn teeth. Prominent overbite 
seems to be the case for specimen BP3 127-34 and possibly for BP2/25, although 
distortion of the skull precludes certainty for the latter. 

As few of the teeth were isolated from their alveoli, examination of the root systems 
was approached through study of radiographic plates. The number of roots for the 
mandibular molars was two for all samples in the series. Maxillary molars, however, 
exhibit greater variation. The distomolar of BP2/20-41 has two roots although the 
other molars adjacent to it are triradical. The first maxillary molars of BP4/8 
are double-rooted. Premolars are single-rooted in the mandibles of both sexes, 
but in the maxilla the first premolar is double for females BPi/6 and BP2/21. These 
double roots are small. 

The direction of the molar roots within their alveoli shows considerable variation. 
The roots of the M^ of BP2/17 are fused and straight. Its neighbour, the M-^, 
shows fusion of only two of its roots — the mesiobuccal and distobuccal. The M^ 
has divergent buccal roots with the radical apices oriented toward each other. 

The mandibular molars of BP2/i7i are divergent, those of the M'l curving toward 
each other. 

Maxillary molars of BP2/20-41 show divergent roots for the distomolar, fused roots 
for the M^, and divergent roots for the remaining molars. The mandibular molars 
all have divergent roots. The medial roots curve distally, and the distal roots are 
straight. 



172 



human skeletal material from ceylon 
Table 8- — Dental Measurements and Indices 









Maxilla 




Mandible 

A 




r 

Range 


Mean 


Range 


Mean 


Third 


R 


Mesio-distal 


8 -5-10 (4) 


9-25 


11-12 (3) 


11-50 


Molar 




Bucco-lingual 


"•5-13 (4) 


11-87 


IO-5-II-5 (3) 


11-00 






Index 


115-00-151 -94 


128-94 


91 -30-100-00 


95-71 




L 


Mesio-distal 


9-10 (4) 


9-37 


10-5-12 (4) 


II -25 






Bucco-lingual 


12-13 (4) 


12-50 


10-5-13 (4) 


11-25 






Index 


120-00—144 '44 


133-80 


87-50-118-18 


100-43 


Second 


R 


Mesio-distal 


9-10-5 (6) 


9-60 


10-5-12 (5) 


11-20 


Molar 




Bucco-lingual 


10-5-13 (6) 


11-75 


9-5-12 (5) 


10-50 






Index 


105 -00-141 -18 


122 -11 


86-36-100-00 


93-67 




L 


Mesio-distal 


9-10 (5) 


9-50 


10-12 (5) 


II-OO 






Bucco-lingual 


10 -5-12 -5 (5) 


11-90 


10-11-5 (5) 


10-60 






Index 


110-53-138-89 


125-44 


91-30-104-54 


96-38 


First 


R 


Mesio-distal 


9-1 1 (6) 


10-17 


10-5-12 (5) 


II-OO 


Molar 




Bucco-longual 


11-12 (6) 


11-50 


IO-II-5 (5) 


10-80 






Index 


109-10-122-22 


I 13-31 


91 -67-104-76 


98-33 




L 


Mesio-distal 


9 -5-1 1 (5) 


10-20 


10-11-5 (5) 


10-80 






Bucco-lingual 


11-12 (5) 


11-70 


10-5-11-5 (5) 


lo-go 






Index 


109- 10-121 05 


1 14 -90 


95-45-105-00 


100-13 


Second 


R 


Mesio-distal 


6-7 (6) 


6-75 


7-7-5 (4) 


7-12 


Premolar 




Bucco-lingual 


9-10 (6) 


9-67 


8-5-9(3) 


8-83 






Index 


128-57-153-85 


143-50 


120-00-128-57 


123-33 




L 


Mesio-distal 


6-5-7 (5) 


6-90 


7-7-5 (4) 


7-25 






Bucco-longual 


9-5-10 (5) 


9-80 


8-8-5 (4) 


8-12 






Index 


I35-7I-I53-85 


142-20 


106-67-121 -43 


112-26 


First 


R 


Mesio-distal 


6-7-5 (5) 


6-80 


7-7-5 (3) 


7-17 


Premolar 




Bucco-lingual 


9-10 (6) 


9-50 


8-9 (3). 


8-33 






Index 


128-57-158-33 


140-42 


114-28-120-00 


116-19 




L 


Mesio-distal 


6-5-7-6(5) 


6-90 


6-5-7-5 (4) 


7-00 






Bucco-lingual 


9-10 (5) 


9-70 


8-8-5 (4) 


8-12 






Index 


128-57-153-85 


140-95 


106-67-123-08 


116-36 


Canine 


R 


Mesio-distal 


6-5-7 (6) 


6-92 


5-5-7 (2) 


6-25 






Labio-lingual 


7-10 (6) 


8-33 


9-5(1) 


9-50 






Index 


100-00-142-86 


120-42 


172-73 


172-73 




L 


Mesio-distal 


6-5-7-5 (4) 


6-75 


5-6-5 (3) 


5-83 






Labio-lingual 


8-8-5 (4) 


8-37 


7-9-5 (3) 


8-00 






Index 


113-33-130-77 


124-49 


I 15 -38-190 00 


140-68 


Lateral 


R 


Mesio-distal 


6-5-7 (2) 


6-75 


6(1) 


6-00 


Incisor 




Labio-lingual 


6-5-7 (2) 


6-75 


6(1) 


6-00 






Index 


92-86-107-69 


100-27 


100-00 


100 • 00 




L 


Mesio-distal 


6-7(4) 


6-62 


5-5(1) 


5-50 






Labio-lingual 


6-5-7-5 (4) 


7-00 


6(1) 


6-00 






Index 


100-00-108-33 


105-79 


109-09 


109-09 


Central 


R 


Mesio-distal 


8-5(1) 


8-50 


. . 




Incisor 




Labio-lingual 
Index 


7(1) 
82-35 


7-00 
82-35 




-• 




L 


Mesio-distal 


9(1) 


9-00 


7(1) 


7-00 






Labio-lingual 


7(1) 


7-00 


6.5 (I) 


6-50 






Index 


77-78 


77-78 


92-86 


92-86 



A'^.B. The small size of the dental series has led the writer to include the data of male and 
female specimens together in the tabulations of Ranges and Means. The numbers in parentheses 
refer to the frequency of observations. 



HUMAN SKELETAL MATERIAL FROM CEYLON 173 

The double roots of the M^ of BP4/8 converge, but are distally oriented. The 
remaining molars of the maxilla diverge and are also distally oriented. 

Among the females, BPi/6 shows three converging roots for the M^ and M^, but 
divergent roots for the M^. The mandibular molars also have divergent roots. The 
M^ has a medial root that is distally oriented. The M^ shows distal curvature for 
both its roots. 

For specimen BP2/21 the radical apices of the maxillary second molar are oriented 
distally, but this trait is less prominent in the other maxillary molars. Mandibular 
roots are divergent and are distally overted. The tips of the roots of the M^ which 
has not completely erupted, are still not fully developed. 

BP2/25 has fused roots for the M— and divergent roots for the Mi, the apices of 
which are curved toward each other. The mesial root shows the greatest curvature. 

The mandibular molar roots of BP3/27-34 are short and distally oriented. The 
LM2 has a functional neck that includes the upper third of its roots. The other 
molars have their roots buried in their alveoli. 

The direction of the premolars is most usually straight. The mandibular pre- 
molars of BP2/21 and BP3/27-34 are curved distally, particularly at the radical apex. 

The pulp cavities vary in size for the molars, but in the maxilla they are universally 
within the cynodont class. Taurodontism is present in the mandible as seen in the 
third molars of BP2/20-41 which is mesotaurodont and in BP2/21 which are hypo- 
taurodont. The second mandibular molar is mesotaurodont in BP2/21 and hypo- 
taurodont in BP2/25. The M^ is mesotaurodont for BP2/21, thus giving to all the 
lower molars of this specimen the taurodont condition. 

Pulp cavities for the other teeth are variable. They are large for canines and 
incisors, but for premolars greater differences are observable. The maxillary 
premolars of BP2/17 have sinuous pulp chambers. Those of BP2/20-41 are broad. 
These canals are straight. BP4/8 has narrow chambers. Those of BP2/21 are 
short for the maxillary premolars but long and broad for the mandibular premolars. 
The largest pulp cavities for premolars are found in the mandible of BP2/25. The 
premolars of the remaining specimens are of moderate size (Pis. 7, 9, 14-15). 

The Biochemical Analysis 

{Table 9) 

Fifteen samples of ground bone from the Balangoda skeletal series were selected 
as suitable for an analysis of their biochemical properties. Of these, two were 
analysed for amino acids by chromatography, three for antigen content by two 
paleoserological techniques, four for nitrogen content, five for the presence of fluorine, 
iron, calcium carbonate and the phosphate P2O5, and six were subjected to radioactive 
methods of dating. Criteria for the selection of these five samples were : (i) the 
observation of osseous tissue in the ground bone sample when subjected to low power 
microscopic examination ; (2) the separation of osseous tissue from the gross in- 
organic content of the sample (concretion, sand, dirt, etc.) after rapid and prolonged 
centrifugation in a saline suspension. 



174 



HUMAN SKELETAL MATERIAL FROM CEYLON 



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HUMAN SKELETAL MATERIAL FROM CEYLON 175 

Chemical Assay (Table lo). In order to reduce the effects of contaminates in 
the ultimate samples of ground bone, the skeletal fragments were cleaned of surface 
dirt by means of a coarse-burred dental drill (Flat Fiss No. 9) and a wire brush. 
Those which were to be subjected to nitrogen (and paleoserological) analysis were 
washed in a solution of 2% absolute ethyl alcohol (denatured) and 98% distilled 
water, then air-dried for 30 minutes and oven-dried for i hour. The diploe was 
extracted from each bone fragment with the use of the coarse-burred drill after an 
orifice for penetration had been cleaned with a finer drilling piece (Flat Fiss No. 6). 
For cranial fragments, a portion of the left parietal i-J" x ih" was removed from the 
brain case with a high-speed circular saw (HSS. 18). The sides of the section were 
cleaned with the fine-burred drill before the diploic tissue was extracted. Large 
bits of bone in each sample were separated from the powdery bone and inorganic 
residue by sifting. The ground bone samples were placed in a mortar where they 
underwent further pulverization. These samples were then deposited in labelled 
and sterilized bottles which were tightly sealed. Securing the sample from the 
diploic tissue has the virtue of reducing the risk of contamination as well as preserving 
the external morphology of the skeletal specimen for further anthropometric study. 
The cranial sections can be reset into the parietals from which they were taken without 
visible evidence of the marks of the saw. The diploe may be substituted with plaster- 
of-Paris in order to prevent the walls of the section from being crushed. It was at 
this level of preparation that the five specimens were selected from the series of nine 
for biochemical analysis. 

The chemical assay of the Balangoda skeletal series was conducted by Mr. G. F. 
Phillips of the Laboratory of the Government Chemist, London. Nitrogen analysis 
was reported by Mr. G. C. Ross of the Central Laboratory, Department of Zoology, 
British Museum (Natural History). The nitrogen content and radioactivity of 
specimen BP2/21 had earlier been reported by Dr. Kenneth P. Oakley (Deraniyagala 
1960a : 97). It should be noted that the results obtained from specimen BP2/25 
were variable as the sample contained a high proportion of sand. 

Amino Acid Chromatography. This analysis of two bone samples from the 
Balangoda skeletal series — BP2/17 and BP3/27-34 — is taken from the report of 
Mr. Ross. 

" The samples were treated with 6N HCL (5 ml. to approx. o-i-o-3 gms.) and 
hydrolysed at boiling pt. under reflux condensation for 16 hours. After suc- 
cessive evaporations to dryness in vacuo over cone. H2SO4 and caustic soda, and 
resolution in distilled water (to remove excess NCI) the filtered hydrolysates 
were subjected to electrolytic desalting to remove excessive inorganic ions 
present. To facilitate the desalting the solutions were diluted to about i in 10. 
After final evaporation to dryness the residue was dissolved in 0-5 ml. water 
prior to chromatography." 

Mr. Ross has noted that the quantities of amino acids present in these two samples 
are extremely small. The limited quantities of material available for examination 
and the short time available for analysis have contributed to the fact that positive 



176 HUMAN SKELETAL MATERIAL FROM CEYLON 

chromatograms could not be completed. Such duplicated chromatograms would 
have contained all suspected known substances run in conjunction and parallel 
with the mixtures of unknown composition. Furthermore, because of the trace 
quantities of nitrogenous material the problem of the elimination of contaminants 
was not investigated as thoroughly as it would have been had more time been 
available. 

" One way chromatograms were prepared on Whatman No. i sheets measuring 
22J" X 18" to give ' runs ' of approx. 18 inches. Each sheet contained base 
line spots of 0-2 ml. of each extract interspersed with 25 spots of known amino- 
acids each spot containing proline for reference Rp values. 

" The runs were made as follows : 

Sheet A. Solvent : Tertiary Butanol : Methyl Ethyl Detone : water : 
diethylamine :: 80 : 80 ; 40 : 8. Ascending 22 hours. 

Sheet B. Solvent : Ethanol : water : Ammonia :: 180 : 10 : 10. Descend- 
ing. Overnight. 

Sheet C. Solvent : n-Butanol : acetic-acid : water :: 120 : 30 : 50. 

Sheet D. Solvent : 80% Phenol : Ethanol : Ammonia :: 150 : 40 : 10, 
Ascending. Overnight. 

Sheet E. Solvent : Methanol : water : Pyridine :: 160 : 40 : 8. Ascending. 
Overnight. 

" Sheets were dried in a cold air stream. 

" Spots were located by dipping through 0-2% Ninhydrin in acetone containing 
2% Pyridine and developing in a warm air stream for | hour then leaving for 
24 hrs. at room temperature. After marking the spots the colours were made 
permanent with alcoholic copper nitrate reagent. 

" Distances of centres of spots were measured from the base-line and (Proline) 
values obtained for all substances except on sheet C where the proline was not 
added (to detect if any in the tests) and sheet B where the proline and substances 
of higher Rf values overran the paper. 

" Results. The listed amino-acids are those probably present as determined 
by single substance spots shown on one or more chromatograms and/or by 
process of elimination from the combinations or substances in ' multiple spots '." 

These results are the following : 

Specimen BP2/17 : (Granular) (Pulverized) 

Palantine Palantine/Tyrisine* 

Glutamic acid Glumatic acid 

Glycine Glycine 

Leucine/Iso Leucine Leucine/Iso Leucine 

Methionine Methionine 

Serine Serine 
Glutamine 

Lysine Crystine/Cysteine 

Taurine 



HUM AN SKELETAL MATERIAL FROM CEYLON 177 

Specimen BP3/27-34 : Palinine Palinine 

Cr Justine /Cysteine Crystine /Cysteine 

Glutamic acid Glutamic acid 

Glycine Glycine 

Leucine/Iso Leucine Leucine/Iso Leucine 

Methionine Methionine 

Ornithine 

Serine /Asparagine Serine 

Taurine Taurine 

Arginine /Asparagine * 
Thrionine? 
Aspartic acid? 

The asterisk (*) following the notation of certain of the above mentioned substances 
indicates that either one or both are present. 

Palaeoserology (Table 10). Bone fragments used as samples for palaeoserologi- 
cal procedures were washed and dried by the methods already described. After 
pulverization of the diploe for each of the three specimens, the samples were weighed 
out into 0-2 gm. amounts and placed in sterilized 75 X 10 mm. tubes which were then 
tightly stoppered. These were reserved for the Standard Absorption Technique. 
Bone samples for the Alcohol Extraction Test were prepared by the same methods, 
but the amount of bone in this second procedure was i-o gm. for two specimens. 
These amounts were placed in two 25 ml. Earlemeyer flasks bearing the appropriate 
label for each of the samples. 

Standard Absorption Technique. This procedure is one based upon the presump- 
tion that a group specific antigen is present in the given sample if antiserum of 
proper concentration is inhibited in its activity with red cells of the appropriate 
group following its prolonged contact with the test sample. It is a technique 
developed originally for dried saliva and muscle tissue samples (Boyd & Boyd 1933, 
1934), and later modified for bone samples (Candela 1936). It is a variation of the 
Standard Inhibition Test for blood group-containing substances. 

Four naturally occurring human Anti-A and four Anti-B sera from individuals of 
known serological constitution were selected for use with each of the three bone 
samples. These anti-sera were recognized as yielding reliable results from previous 
testings with controls of known blood groups. In setting up a titration series, 
buffered saline, to which had been added 1% sodium azide as a preservative, was 
used as a dilutant. The determination of the titre for the anti-sera was established 
as the third or fourth dilution from the end-point for any one serum as judged from 
microscopic examination of a smear on a glass slide. A panel of anti-sera was 
regarded as preferable to pooled anti-sera by this investigator. The volumetric 
unit was used for all titrations in this analysis since it allows for greater accuracy 
than the more commonly used drop technique. To the 0-2 gm. of ground bone 
deposited in each of the labelled tubes was added 0-5 cc. of neat serum of the appro- 
priate type. After these contents were mixed, the tubes were placed in the refrigera- 



lyS HUMAN SKELETAL MATERIAL FROM CEYLON 

tor for an incubation period of 24 hours. At the termination of this period of time, 
the tubes were centrifuged for 4 minutes at moderate speed. The supernatant 
was then drawn from each tube by means of a Pasteur pipette with its tip encased 
in sterihzed cotton wool. The titration for the sample series and a control series 
was then conducted. Both series contained equal volumes of saline. In the sample 
series titration was carried out with the supernatant, in the control series with the 
neat sera. To each were added equal volumetric units of 1% suspension in saline 
of red cells of the appropriate type. The sample and control series were then 
incubated for 2 hours at room temperature. At the termination of this period smears 
from each of the tubes were placed on glass slides and interpreted under microscopic 
examination. Three tests were conducted over three 24-hour intervals. 

Results are based upon the presence of agglutination in one or both of the anti-sera 
or the absence of agglutination in both anti-sera for any given sample. If agglutina- 
tion occurs in both the Anti-A and Anti-B sera, as a result of the introduction of red 
cells of appropriate type, the blood group is assumed to be O. Likewise if agglutina- 
tion is absent from both anti-sera, AB type blood is suggested. Type A blood is 
marked by the presence of agglutination in the Anti-B sera but absence of agglutina- 
tion in Anti-A sera. Type B blood is marked by a reversal of this situation. 

Alcohol Extraction Test. This technique is devised to reduce the problem of 
non-specific absorption and thus allow for the correct identification of blood type AB 
(Gray 1953, 1958). It was applied to the bone samples X and Z noted in Table 10. 
Five different anti-sera were selected for the test. 

In each of two 25 ml. Earlemeyer flasks, which were labelled x and z for the two 
samples being tested, was deposited i-o gm. of ground bone of the appropriate 
specimen. Both flasks were then filled with 25 ml. of absolute ethyl alcohol (de- 
natured). Each preparation was heated for a period of 2 hours at 78 degrees C. 
The alcohol extracts, which contained the group substance, were filtered off and placed 
in sterilized beakers. To the bone residues were next added 15 ml. of absolute ethyl 
alcohol and each preparation was heated as above. After filtering this second 
quantity of alcohol extract, it was added to that extract initially collected in the 
beakers. The two extracts were placed over a steam bath until the lipid material 
began to come out of solution. The extracts were removed from the heat to re- 
dissolve the precipitate. The contents of the beakers weie then deposited in sterilized 
bottles which were tightly stoppered. These extracts are the stock antigens. The 
antigen suspensions were prepared by first placing i-o ml. of each stock antigen into 
test tubes containing 5-0 ml. of buffered saline. Milky-appearing emulsions resulted. 
These were used for the determination of the presence of blood group antigens by 
first adding 0-5 ml. of each antigen suspension to 0-5 ml. of Anti-A and Anti-B sera 
whose titres had already been established. These were mixed in 10 x 75 mm. tubes 
and incubated at room temperature for 2 hours. At the end of the incubation period, 
the tubes were centrifuged for 4 minutes at high speed. Then 0-5 ml. of supernatant 
was removed from each tube and placed in a sterilized tube by the same technique 
described for specimens in the Standard Absorption Technique. A titration for 
both samples and controls was arranged. To equal volumes of saline, placed in all 



HUMAN SKELETAL MATERIAL FROM CEYLON 179 

but the first tubes of the panel, were titrated the neat sera with the controls and the 
supernatant with the sample series. Each tube received equal volumes of a 1% 
suspension in saline of red cells of appropriate type. Samples and controls were then 
incubated for two hours at room temperature. 

After incubation the contents of each tube were read under the microscope and the 
results were recorded. This technique was limited to a single test. 

Interpretation of these results is based upon the following conditions : 

1. The known type A extract suspension should lower the titre of Anti-A by 
3 or more tubes. 

2. The known type B extract suspension should lower the titre of Anti-B by 3 or 
more tubes. 

3. The known type O extract suspension should not lower the titre of either 
Anti-A or Anti-B by more than i tube. 

4. If the test extract suspension lowers the titre of Anti-A by 3 tubes or more 
then the sample is type A. 

5. If the test extract suspension lowers the titre of Anti-B by 3 tubes or more then 
the sample is type B. 

6. If the test extract suspension lowers the titre of both Anti-A and Anti-B by 
3 tubes or more then the sample is type AB. 

7. If the test extract suspension lowers neither the titre of Anti-A nor Anti-B by 
more than i tube then the sample is type O or else the antigens in the sample have 
deteriorated to the point where conclusive typing is impossible. 

Radioactivity Methods of Dating. Radiocarbon analyses of human bone and 
charcoal from Bellan Bandi Palassa have yielded results that confirm Deraniyagala's 
(ig43a : 112 : 1945) contention that the Balangoda Culture persisted until com- 
paratively recent times and existed for some time side by side with ferrolithic 
cultures. 

Two samples of charcoal were tested by Isotopes Incorporated in 1956. Sample 
394K was taken from 2' 8" below the surface and i' above the limestone bed rock in 
Square Ci. This was the locus of skeletal specimen BP3/i5a. The charcoal lay 
3' north of the skull of this specimen. Its dating is 508 ± 150 years B.P. or 1448 
A.D. i 150 years. Sample 394L was taken from 2' 6" to 3' below the surface and 
almost resting upon the bed rock in Square Bi. This was within the vicinity of 
skeletal specimen BP2/20. Its dating is 2070 ± 200 years B.P. or 114 B.C. ± 200 
years (Deraniyagala 1958a : 259). 

The radioactivity of the bone sample was assessed in the laboratory of the Anthro- 
pology Sub-Department, British Museum (Natural History), in 1959. This was a 
sample of tibia from specimen BP2/21. This skeleton was found in Square C4 
some 3' below the surface and 6" above the limestone. Oakley's report to Deraniya- 
gala stated : 

" In answer to your previous letter, I can tell you that the radioactivity of the 
human leg bone from the Ballanbandi Palassa site indicated 0. 6 p. p.m., eU3o8. 
This is consistent with the specimen being ancient — say, having an antiquity 



i8o HUMAN SKELETAL MATERIAL FROM CEYLON 

measured in millenia rather than in centuries. Chemical analysis shows 1-3% 
nitrogen which suggests in the circumstances of its occurrence, that the bone is 
Post-Pleistocene " (Deraniyagala 1960a : 97). 

In 1962 osseous specimens from the Balangoda series were assayed by Mrs. 
Elizabeth Gardiner in the same Department. In a personal communication con- 
cerning the results of the assa}^ Oakley reports : 

" A sample of the ribs of skeleton BP3/27-34 (D) was assayed completely (For 
three hours), and gave the result (calculated as eU3o8 p. p.m.) : 10 ± 2. The 
remaining Balangoda samples were assayed for half an hour each and gave 
readings which when averaged amounted to approximately the same figure. 
As the order of magnitude is low I do not think that anything would be gained 
by assaying all the samples completely." 

Oakley notes that all of the samples proved to have a uranium content that was 
fairly uniform. 

Table 10 
Results of the Biochemical Analysis 

Chemical Assay : 





Skeletal 


Ratio 












specimen 


X 100 












Number %F %V^O, F 


:P205 


°/ 


;,Fe %CaC03 


%N 






BP2/21 








1-3 






BP3/27-34 o-6o 23-1 


2-6 




2-5 17-1 


0-05 






BP2/i7g 0-38 12-5 


30 




2-8 21-6 


0-078 






BP2/17 0-32 i8-o 


1-8 




5-0 23-7 


o-oi6 






BP2/21 0-58 21-7 


2-7 




1-5 19-5 








BP2/25 0-15 7-0 


2-0 




3-0 10-5 






Palaeoserology : 












Standard 


Absorption Technique 












Test No. 


I 


2 






3 




Anfi- 


A 


A 






A 




Sera 


1 \ 1 
81-02 90-11 B-25 R-58 81-02 


90-11 


R 


-25 R-58 81- 


-02 90-11 R- 


-25 R- 


Sample 














X 


A 

















Y 


A 





A 











Z 


AAA? A 


A 


A 


A A 


A A 


A 



Alcohol Extraction Test 

Anti- 
Sera Gr-02 42-46 51-59 
Sample 

X A A AB 

Z AB A AB 



Of these five techniques of analysis of the biochemical nature of the osseous material 
from Bellan Bandi Palassa, four are related to the problems of chemical dating : 



HUMAN SKELETAL MATERIAL FROM CEYLON i8i 

the palaeoserological analysis stands apart from these considerations save in so far 
as the identification of antigens can verify the presence of organic elements in the 
bone samples. Since the accumulation or depletion of substances in ancient bone is 
ultimately dependent upon the characteristics of the environment in which it has 
been deposited, and since these environments may alter due to redisposition of the 
bone and/or disturbances of the deposit, it is not surprising that the results of the 
biochemical analysis of these Balangodese specimens exhibit some variations. Not 
all of the skeletal remains from Bellan Bandi Palassa occurred at the same level. 
Thus specimen BP3/27-34 rested only 3" above the limestone bed rock and its 
contact with the hard cr3'stalhne limestone caused the bones to become coated with a 
yellowish concretion, a very different micro-environment from that of specimen 
BP3/i5a, from which level one of the charcoal specimens was taken for radiocarbon 
dating. 

The presence of nitrogen in bone indicates protein and protein derivatives. Nitro- 
gen quantities of the order of hundredths of 1% have no significance since the method 
of analysis becomes progressively less reliable when working with lower quantities 
(Barber 1939, Cook i960 : 229, Pin 1950). Therefore the value obtained from the 
sample for specimen BP2/21 is more significant than the values obtained for the 
other specimens whose nitrogen content was examined. This discrepancy may be 
due to the fact that cancellous osseous tissue was used in these tests instead of the 
compact bone which offers more reliable analytic results. Oakley (1949, 1953, 1955) 
finds that the nitrogen values are reliable as a cross-check to fluorine analysis, 
particularly since the results from both kinds of tests may show radical differences 
for a single specimen. But calcite has a sealing effect on bone, and at Bellan Bandi 
Palassa, where the skeletal specimens lie at different levels above the limestone bed, 
this phenomenon cannot be excluded from a consideration of the results of tests. 
Temperature and moisture are other factors influencing the chemical nature of bone, 
and the evidence for believing that climatic changes have affected particular biotic 
modifications in the region around Bellan Bandi Palassa is considerable. Because 
of these factors of alteration and contamination, Oakley (1953 : 52) has the phosphate 
content of each sample checked and he regards the percentage fluorine/phosphate 
ratio as the best standard of comparison in fluorine dating. He has found that the 
average fluorine content of Lower Pleistocene bones is circa 2-o% ; of Mid-Pleistocene 
bones, circa i'5% ; and of Upper Pleistocene bones, circa 0*5%. The percentages 
for the Bellan Bandi Palassa specimens fall within this latter category, a situation 
confirmed by the radiocarbon data for the site which marks the age of the site as 
post-Pleistocene. 

The development of chromatographic techniques has expanded the data of protein 
analysis to include not only nitrogenous components but also the amino acids (Abelson 
1954, 1955, Ezra & Cook 1957). Bones of a woolly rhinoceros from an Upper Pleisto- 
cene deposit of the Lloyds site, London, contained traces of glutamic acid, B-alanine, 
proline, OH-proline, leucine, valine, glycine, aspartic acid and arginine. (The writer 
is indebted to Dr. K. P. Oakley and Mr. A. E. Rixon of the British Museum (Natural 
History) for the notes of this research carried out in 1956. For illustration of the 



i82 HUMAN SKELETAL MATERIAL FROM CEYLON 

Lloyds bone chromatogram see Oakley 1963). Those amino acids present in the 
Balangoda series but absent in this Pleistocene sample are taurine, serine, methionine, 
ornithine, cystine, glutamine, theonine (?) and tyrosine (?). The presence of different 
percentages by weight of amino acids for blood group substances may be explained 
by variations in the preparation of the serological sample and hence bear no relation 
to immunological specificity (Kabat 1956 : 149-151). However, should certain 
blood group substances be demonstrated as having characteristic components of 
amino acids, the findings of palaeoserology would become pertinent to problems of 
chemical dating. The nature and quantity of protein decomposition products in 
ancient bone might well be utilized as a method or technique of chemical dating which 
Cook (i960 : 232) suggests would be superior to the analyses of total organic matter 
or nitrogen content. 

The problem of contaminants cannot be minimized for any of these biochemical 
tests but their effect, if present, is most serious in the establishment of the nature of 
the antigen content. It is impossible at present to detect the presence of foreign 
substances in the sample which would bias the results of analysis, but by running the 
sample through several tests and subjecting it to different techniques of analysis 
the researcher is encouraged in verifying a certain percentage of his results. Thus, 
the behaviour of sample Y in the Standard Absorption Technique cautions reservation 
of opinion since it may have undergone a non-specific absorption with the Anti-A 
sera. The Z sample is more reliable, its consistent type A reaction being hindered 
only at one place in the test due to lysis of the Anti-B serum. Of the four Anti-A 
sera used, the B sample is somewhat erratic in its behaviour and its unsuitability 
for testing with these bone samples is suggested. The results of the Alcohol Extrac- 
tion Test cannot be accepted as being as reliable as the data derived from the initial 
test for antigen identification. Reactions are all very weak. The presence of type A 
antigen for sample Z is confirmed by both tests and the Alcohol Extraction Test 
suggests that this may be linked with the B antigen, thus indicating for specimen Z 
an AB blood type. The occurrence of the A gene in the Balangodese series is of 
particular interest since this gene has a very low frequency among the Veddas and 
its presence in this population in the past has been doubted by some investigators 
(Wickremasinghe, Ikin & Mourant 1963). 

The presence of pottery at the upper portions of the Bellan Bandi Palassa midden 
and its absence in the lower portions has led Deraniyagala (1958a : 258) to conclude 
that the site had been occupied either continuously or periodically over a period of 
sufficient duration to allow the inhabitants to pass through both phases of his 
Balangoda culture. Furthermore he had suggested a correlation of cultural strati- 
graphy with the physical type of the skeletal specimens found in the site : the robust 
male specimen BP2/17 appears to him as " Australoid " while the specimens from 
the upper pottery-bearing levels reveal " Negroid " phenotypic traits. Regardless 
of what racial features one may perceive in the series, the radiocarbon methods of 
dating affirm that these specimens were relatively contemporary in time. In the 
opinion of the present writer it seems improbable that the specimens from Bellan 
Bandi Palassa constitute a stratigraphical progression of phenotypes representative 



HUMAN SKELETAL MATERIAL FROM CEYLON 183 

of earlier and later phases of a pre-metal using culture, nor do the artifacts found in 
association with the burials suggest a cultural succession : the phenotj^pic variation 
which the series displays is what can be expected from normal sexual dimorphism 
and morphological variation across sexual lines, and the question of cultural sequences 
at the site cannot be answered withoiit confirming evidence from further archaeologi- 
cal investigation of other Ceylonese sites. If the osseous specimens were not all 
members of the same community, the dating of the bones by radiocarbon methods 
indicates that they were not separated by many generations. 

Ill COMPARATIVE ANALYSIS 

The Nature of the Comparative Data Used in the Determination of the Biological 
Affinities of the Balangodese with Other Populations 

In compiling the comparative data of Tables it and 12 the writer has selected 
several contemporary and prehistoric populations and individual specimens from the 
Indian-Southeast Asiatic--Oceanic area. The arrangement of the prehistoric series 
in the Tables corresponds to their chronological relationship to the modern popula- 
tions listed. Of these, the writer has examined at first hand the Vedda series and 
the specimens from Brahmagiri, Adichanallur, Mohenjo-daro, Nal, Langhnaj, 
Bayana, Sialkot and Niah, and casts of Talgai, Keilor and Wadjak. Comparisons 
have also been made with the original specimens from such Indian prehistoric sites 
as Nevasa, Chandoli, Maski, Piklihal, Harappa, Lothal, Ruangarh and Raigir, whose 
metrical values are not included in Tables 11 and 12. On the basis of this extensive 
comparative analysis it is the thesis of the writer that of the living populations of 
southern Asia the Veddas of Ceylon most closely resemble the Balangodese in their 
physical morphology. It must be emphasized that the fragmentary nature of the 
Balangodese skeletal remains limits their significance in a purely metrical comparative 
analysis with larger and more complete osteological specimens. Hence the data of 
the tables are less helpful than a morphological comparison in affording the reader a 
clear idea of the phenotypic similarities of the prehistoric inhabitants of Ceylon. The 
Crown Indices of Table 12 have been calculated with the formula 

Bucco-Lingual Diameter 
Mesio-Distal Diameter 

It should also be noted that the series of five specimens from Brahmagiri are 
regarded as males although another investigator has regarded specimen IE as female 
(Sarkar i960). 

The Vedda osteological specimens employed in this analysis were examined by 
the writer from collections from the following institutions : Sub-Department of 
Anthropolog}^ British Museum (Natural History), London ; Royal College of 
Surgeons, London ; Anatomical Museum, University College, London ; Museum of 
Comiparative Anatomy, University of Oxford ; Museum of Human Anatomy. 
University of Cambridge. The comparative study was supplemented by the com- 
prehensive work on the physical anthropology of the Veddas by Hill (1941). In his 
study of this population Hill makes a distinction between specimens that he 

GEOL. 11,4 17 



i84 HUMAN SKELETAL MATERIAL FROM CEYLON 

regards as " pure Veddalis " and those of " half-breed Veddah, portraying . . . 
certain distinctive Veddah features upon a basically different structure." (Hill 1941 : 
107, 124-125). This dichotomy is a subject of discussion elsewhere in this paper, 
but the terminology that Hill employs is repeated here when comparing Balangodese 
specimens with the Veddas of his " Average " series and " Selected Average " 
series. Reference was made as well to Stoudt's (1961) anthropometric survey of 
living ethnic groups in Ceylon, a study based in part on data collected by the late 
J. R. de la Haule Marett. 



A Comparison ivith the Veddas 

The Calvarium. The Veddas and Balangodese are dolichocranic. Mean indices 
are 71-24 for Vedda males and 72-72 for Vedda females, with a " Selected Average " 
of 71-57 for specimens of known Vedda ancestry. The Vedda female has a Cranial 
Length-Breadth Index slightly greater than that of the male. Hill notes that the 
discarding of the indices of Vedda specimens of doubtful purity makes little difference 
in this Index, the average for all Vedda crania being 71-60. The Length-Height 
Index based on Basion-Bregma Height averages 74-64 for both male and female 
Veddas, thus placing them at the upper limit of the orthocranic class. However a 
number of Vedda crania which Hill regards as belonging to pure-blooded 
individuals have Basion-Bregma Height-Length Indices that place them in the 
hypsicranic group. When the Vedda series is examined with respect to the Auricular 
Height-Length Index, the male mean of 61-00 falls within the orthocranic category, 
but the females are hypsicranic with a mean index of 63-50. Here is a situation that 
is not paralleled in the Balangoda series where the male is hypsicranic and the 
female chamaecranic. There is closer agreement between the two populations in 
terms of the relation of cranial height to cranial breadth since the acrocranic condition 
which is common to Veddas of both sexes, is also apparent in the Balangodese male. 
The low Cranial Height-Breadth Index of the Balangodese female stands outside the 
mean values of 105-5 for Vedda males and 103-9 for Vedda females. 

The " Selected Average " of cranial capacities for male and female Veddas are 
1250 cc. and 1166 cc. respectively. Although the addition of values for crania of 
dubious ancestry does not raise these mean values to a marked degree, the values for 
unmixed Tamils and Sinhalese are significantly higher than those for unmixed Veddas. 
These low values include within their range the estimated cranial capacity for the 
Balangoda female BP3/27-34, but the large-headed male, BP2/17, falls well outside 
the range. A further comparison of cranial size is offered by the measurement of 
the Maximum Circumference of the vault where the values for the Veddas are close 
to those of the Balangoda females, but from which again the male BP2/17 stands 
apart. 

Vedda cranial bones, which are uniformly thick and heavy, particularly in their 
posterior regions, are regarded by Hill as belonging to individuals of doubtful purity 
and it is these which most closely approximate a condition evident in the Balangodese 
crania. The latter series lacks the pathological thinning peculiar to the parietals of 



HUMAN SKELETAL MATERIAL FROM CEYLON 185 

Vedda crania of the classic type. Both series exhibit little or no sexual differentiation 
in cranial thickness. 

The Vedda crania observed by the present writer were rhomboid although the 
crania of males with well developed supraorbital ridges approached a sphenoid 
cranial conformation. Supraorbital ridges are well developed in both Veddas and 
Balangodese. and the very pronounced ridges of BP2/17 are occasionally found among 
the Veddas as well. Confluence of ridges at glabella is present in half of the Vedda 
series and in both sexes, but among the Balangodese females the brow ridge form is 
median. A low vertical forehead with flattened parietals and a posteriorly com- 
pressed occiput is the most typical Vedda cranial contour, and it is this form which 
also is characteristic of the Balangodese females. The medium receding forehead of 
BP2/17 has been observed among Veddas where it is accompanied by some lateral 
compression of the frontal. The presence of a median boss and frontal eminences 
are frequent in the crania of both series. The Balangodese females approach that 
degree of frontal constriction common to Vedda crania, but the frontal of BP2/17 
is considerably broader than that of most Vedda males. Supraorbital foramina are 
prominent in both series. 

Slight keeling of the parietal region is both a Vedda and Balangodese trait. Pos- 
terior to vertex the vaults of crania from both series undergo a gradual slope to 
lambda, at which point a sudden change of direction brings the curve vertically 
downward to the basi-occiput region. The flattening of regions anterior-superior 
and posterior-inferior to the parietal eminences contributes to the prominence of 
the bosses in the vaults of both series. Differences appear in the occipital portion of 



FRANKFORT HORIZON 



Fig. 5. Left lateral contours of one Balangodese (BP3 127-34 ), one Vedda (1949. 

12.7.2 ), a compound tracing of eighty-two male Australians ("Wagner 1937 

), a compound tracing of one hundred and twenty-four male New Guineans 

(Hambly, 1940 -.-.-). 



1 86 



HUMAN SKELETAL MATERIAL FROM CEYLON 




Fig. 6. Frontal contours of one Balangodese (BP3/27-34 — ^— ^— ), one Vedda (1949. 

12.7.2 ), a compound tracing of eighty-two male Australians (Wagner 1937 

), a compound tracing of one hundred and twenty-four male New Guineans 



(Hambly, 1940 



the cranial profile. The musculature of the nuchal crests is weakly developed among 
the Veddas, a condition reflected in the Balangodese females but not in the robust 
male BP2/17 (Text -figs. 5-6). 

The temporal muscles occupy extensive areas of the crania in both populations. 
The temporal lines rise high along the vault of the Vedda skulls. They are less 
prominent on the parietals of the Balangodese. The mastoid process is generally 
small among the Veddas, but several skulls of Hill's series possess large processes 
like that of BP2/17, and these generally belong to males of dubious ancestry. The 
tympanic plate is thick in Veddas and Balangodese. The oval form of the Vedda 
auditory meatus contrasts with the elliptical and round conformations of this struc- 
ture in the fossil population. 

Like the Balangodese, the Vedda cranial vault is characterized by suture patterns 
of simple design with some slight complexity apparent at peripheral margins. 
Metopism, which is uncommon in Veddas, is absent in the Balangodese specimens. 
However the former series exhibits a high frequency of Wormian bones, especially 
at lambda and pterion. Sutural patterns are complex in this area in Balangodese 
skulls but Wormian ramifications are absent (Text-fig. 7). 

Hill regards the typical Vedda face as euryprosopic for males and hypereuryproso- 
pic for females, the mean index of his " Selected Series " being 84-14. The males 
average 85-44, the females 79-73. However, for the living individuals that Hill 
examined, the average Total Facial Index is within the leptoprosopic category, the 
class to which the Balangoda female BP3/27-34 is also assignable. Concerning this 
problem he writes : " This feature is evidently very variable among the existing 
Veddah population. Combining this with the fact that previous writers have 



HUMAN SKELETAL MATERIAL FROM CEYLON 



187 



— 







195 _ 






190 _ 




• 


185 _ 


, 




180 _ 


• ■ . • 


■ •©. 


175 _ 




'■ © • 


170 _ 


. « «. . 


■•• 


165 _ 


". 


' 


160 _ 


' 




155 _ 














1 1 1 1 1 


1 1 1 1 1 




Cranial Inde 



Fig. 7. Scatter diagram and frequency polygon of cranial measurements and indices of 
three Balangodese {BP2/17, BP2/21, BP3/27-34) and sixty-two Veddas (Hill 1941). 
Males are represented by a dot (.) ; females by a cross (x). 



placed the Veddahs in the chamaeprosopic class, and also that the index based on 
osteometry is well within the euryprosopic category ... it can only be inferred that 
the results . . . are the effect of recent miscegenation with the leptoprosopic 
neighboring peoples " (Hill 1941 : 74). The leptene upper face of the same Balango- 
dese female stands in contrast to the euryene value of 49-00 for typical Vedda speci- 
mens. Closer agreement of Vedda and Balangodese indices is apparent in the 
chamaerrine nasal forms of both groups. The average for Veddas of the " Selected 
Series" is 55-80 and the male and female mean values are 54-24 and 57-30, res- 
pectively. The Orbital Index is 74-00 for Vedda males and 73-70 for Vedda females, 
hence both come within the chamaeconch class, a condition found in the female 
Balangodese BP3/27-34, but not in the male BP2/17 whose orbits are hypsiconch. 
The External Palatal Indices are brachystaphaline, but the indices for the Balango- 
dese are considerably higher than the mean index of 96-50 of the Veddas of the 



i88 HUMAN SKELETAL MATERIAL FROM CEYLON 

" Selected Series ". The Vedda males have an average External Palatal Index of 
97-06, the females are even lower — 94-30. 

Like the Balangodese, the Vedda face owes its prognathism to the projection of 
the total maxillary or subnasal region, although prognathism of the latter sort is 
characteristic of a number of Vedda specimens. Hill finds that alveolar 
prognathism is more common among the Vedda crania of earlier collections : sub- 
nasal prognathism is more often found among recent specimens. The average 
Vedda Gnathic Index is 96-70 which places it within the orthognathic category, and 
in the "Selected Series" this index drops to 94-50. Males have a mean index of 
95-04, females of 93-37. Facial angles of Vedda crania show that almost all values 
fall within the prognathous category if von Camper's method is used, but when 
using Martin's method almost all the faces are orthognathous. 

Both Balangodese and Veddas have square or quadrilateral orbits. Further 
comparisons of the orbit are prohibited by the imperfect condition or the absence 
of the orbital walls in the fossil specimens. Malars are larger in the latter series, 
and the temporal process of the zygomatic and the zygomatic arch is less attenuated 
in Balangodese. The anatomical aberrations that Hill notes as common to the 
region for Vedda crania — a prominent marginal process, a maxillary-zygomatic 
notch — are not present in the fossil series. Malar projection is variable for both 
populations. The nasal aperture is wide, short, and piriform for the two series. 
The inferior nasal margins are variable in form, the oxycraspedote and orygmocraspe- 
dote conditions being represented in equal frequency for both series. The absence 
of nasal bones in the Balangoda series precludes further comparison of this region. 
The forms of the glabella of BP2/17 and BP3/27-34 infer a deep nasal notch for these 
specimens such as is foimd among the Vedda. The anterior nasal spine is large 





Vedda Specimen 1949 12.7.2 



Balangodese Specimen BP 3 27 



Fig. 8. Dioptrographic drawings. Frontal aspects. 



HUMAN SKELETAL MATERIAL FROM CEYLON 



189 





Fig. 9. Dioptrographic drawings. Right lateral aspects. 

among the Balangodese, but its size is variable in Vedda crania. It is lacking in those 
specimens that have orygmocraspedotj^ The alveolar borders of the palates of the 
Balangoda series are more greatly divergent than those of the Vedda series although 
both series may be described as having the parabolic and elliptical palatal conforma- 
tion. In size and depth the palates of the fossil series attain the higher values 
(Text-figs. 8, 9). 

The Mandible. The Mandibular Index of the lower jaws of Veddas falls within 
the dolichostenomandibular category with an average Index of 8o-ii. This is within 
the range represented by the fossil series. The mean Fronto-Gonial Index of the 
Vedda series is 97-11, and the two indices available for the Balangodese fall on either 
side of this value. The average Zygo-Gonial Index for the Veddas — 135-79 — is 
considerably higher than that obtained for the Balangodese, and this reflects the 
greater bizygomatic diameter of the latter population. 

A comparison of the gross sizes of the mandibles in the two series shows close 
similarity, but those of the Balangodese are marked with greater muscularity and 
an increase in the thickness of the corpus, particularly in the symphysial region. The 
chin is prominent among the latter : Veddas have moderate^ developed mental 
protuberances. In both series the chins are most commonly of the median type. 
Genial tubercles are feebly developed in the Vedda mandibles, and not all Balangodese 
mandibles have prominent genial tubercles, viz. BP2/i7i and BP3 127-34. The 
digastric fossae, which are prominent in some Vedda specimens, are variable in 
development in the fossil series. In both series the corpus tapers posteriorly from 
the symphysis, thus giving a gracile appearance to this portion of the jaw. The 
exception to this is the Balangodese male, BP2/i7i whose corpus is massive through- 
out its length. Mental foramina are large for most Vedda specimens, but their size 
can be matched in only half of the individuals in the Balangoda series. The position 
of these foramina is the same in both series. 

The rami form a wide angle with the corpora in the mandibles of both series. It 
is broad and short in the Vedda series, and broad and somewhat more elevated in 
the Balangoda series. In the fossils the gonia are frequently everted, but this is 



I90 HUMAN SKELETAL MATERIAL FROM CEYLON 

rare among Vedda specimens where the muscular development of the pterygoid 
attachments are reduced. The sigmoid notch is broader and shallower among 
Veddas. The condylar neck is reduced in both series, and a mandibular head of 
small dimensions is the most common condition. As with the Veddas, the lingula 
of the Balangodese mandibles is reduced in size, although strongly marked in the 
male BP2/20-41. Alveolar prognathism is most pronounced in the fossil series. 

The morphological features that characterize the mandible of the Balangoda 
series, namely the deptli of tlie corpus, the greater height of the ramus, and the 
presence of mandibular alveolar prognathism, are traits that Hill finds most frequently 
represented among Veddas of doubtful purity, i.e., individuals with longer and 
narrower faces than those of the classic Veddas with their broad chamaeprosopic 
faces accompanied by short, narrow mandibles. This situation parallels the con- 
ditions noted above for the cranium where the Balangodese reflect some morphological 
features not present in Veddas that Hill regards of pure type. 

The Skeleton of the Trunk. Comparisons of the vertebral columns of Veddas and 
Balangodese reveal striking similarities. The cervical vertebrae are very small in 
the absolute sense and considering the evidence for regarding the stature of the 
Balangodese as taller than that of the Veddas, the sizes of their vertebral elements 
seem disproportionately small. Neural foramina are triangular, and transverse 
processes are short within both groups. However, the bodies are round rather than 
cordiform among the Veddas. The atlases in both series are delicate with sinuous 
posterior arches. The thoracic vertebrae described for the Balangodese fall within 
the range of normal features common to the Veddas. The lumbar vertebrae of the 
fossil series are uniquely robust and have a greater mass than the lumbar segments 
of the recent series. The Balangoda specimen BP2/17 has a Lumbar Index that 
falls within the koilorachic group. While this index is matched by several Vedda 
specimens, the average index of the latter is orthorachic. Broad truncated spines 
are characteristic of the lumbar vertebrae of both series. The instability of the 
lumbar and sacral regions and of the thoraco-lumbar and lumbo-sacral junctions, 
which is a characteristic of the Vedda spine cannot be ascertained for the Balangodese 
due to the paucity of bones from this region. 

The ribs of the Balangodese and Vedda specimens have a thickness ranging from 
3 mm. to 6 mm. with maximum expansion at the sternal end for males of both series. 
The first rib has a small head and a tapered neck. Its grooves for the subclavian 
vessels are apparent. Lower ribs have more massive heads. The degree of individual 
variation among the Veddas for this part of the thorax cannot be established for the 
Balangodese, since their ribs and vertebrae are fragmentary and present for only 
two specimens. 

The sternal corpus of BP2/17 approaches the Vedda form in terms of its slight 
curvature, short and broad dimensions, and its number of segments for sternal rib 
articulations. 

The Skeleton of the Upper Extremity. Except for its large size, the clavicle of 
BP2/17 closely resembles the clavicles of Vedda specimens, as does also the smaller 
clavicle of BP3/27-34. Similarities are found in the oval form of the sternal head. 



H U M A N S K E L E T A L M A T E R I A L F R O M C E Y L O N 191 

the moderate to slight degree of flattening of the acromial head, and a general 
gracilitv of the form of the shaft. The clavicle of the female Balangodese specimen 
differs from the Vedda only with regard to its greater muscularity, and the presence 
of pronounced grooves for the subclavian vessels. Clavicular curvature is poorly 
developed in the females of both series. 

There is some difference in the scapulae of the two series, that of the male BP2/17 
being larger and more robust than the scapulae of the Balangoda female and the 
majority of other scapulae of the Vedda series. Other differences are noted in the 
longer and more rectangular form of the acromion process of BP2/17, its shorter 
and narrower fossa supraspinata, and the greater curvature and length of its corocoid 
process. In the Vedda series, the acromion is usually triangular in form, the fossa 
supraspinata tends to be broad, and the corocoid process is short, massive, and 
flattened from its anterior to posterior aspects. The scapula of the female BP3/27-34 
more closely approximates the Vedda norm, as does that of BP2/17, in all other res- 
pects than those noted above. There is a pronounced obliquity of the spino-vertebral 
angle. The superior border lies in the horizontal plane and the supra-scapular notch 
is absent. This notch is absent in half of the scapulae of the Vedda series. The 
axillary borders of scapulae in both series show an increased thickening in the 
direction of the glenoid fossae. The latter structure is broadest at its inferior portion, 
and its angle to the body of the scapula is slightly elevated and lateral. 

The Balangodese and Vedda humeri are alike in their relatively long diaphj'sial 
lengths and small extremities. Their shafts are straight, but the sigmoid curvature 
that Hill notes for his Vedda series is not apparent among the Balangodese. Due 
to the poor condition of the humeri of the latter group their angle of torsion could 
not be estimated. Like the humerus of the Veddas, that of the Balangodese shows 
a prominent bicipital groove. The high incidence of the perforation of the olecranon 
fossa has been found to occur in only 25-60 per cent of the series examined by Hill 
and this feature is absent in the Balangodese. 

The trait that most strikingly sets apart the Balangodese radius from that of 
the Vedda is its greater muscular development. The Vedda radius is characterized 
by an extensive tuberosity, rounded borders, and slender form. Both series, however, 
present a pronounced hollowing of the volar aspect of the shaft for the reception of 
the flexor pollicis longus. The styloid process is prominent in both groups, save in 
the case of BP3/27-34 where its size is reduced. A lateral curvature of the shaft is 
more common in the Vedda radius; the shafts are straight or, as in the case of the 
female specimens, curvature is in an anterior-posterior direction. In both series 
the head is small, and the neck is narrow and elongated. The ulnar notch is well 
marked and extensive. 

The curvature of the ulna is considerably less prominent among Balangodese than 
among Veddas, and the muscular attachments are more obvious among the former. 
Where the interosseous space is large for the Balangoda series, it is a result of radial 
rather than ulnar curvature. 

The bones of the hand of the immature specimen BP2/21 show certain features 
that resemble the Vedda hand. In gross size the carpals and metacarpals lie at the 



192 HUMAN SKELETAL MATERIAL FROM CEYLON 

upper limit of the range for the Veddas. The total digital formula for the Vedda 
hand is III > II > IV > V > I ; the metacarpal formula for the Balangodese is 
II > III > IV > V(?) > I. The metacarpals of both series show curvature in 
the dorso-ventral plane, but Hill does not regard this as a feature unique among 
the Veddas, since it is found as well among other Ceylonese groups. The shapes 
of the carpal bones which are peculiar to the Veddas are also approximated in the 
hand of the Balangodese, viz. the dumbbell conformation of the scaphoid as a result 
of central narrowness, the cuboidal form of the capitate, and the well-defined ridging 
of the trapezium. 

The Skeleton of the Lower Extremity. While the Vedda femora are platymeric with 
an average index of 78-50 for males and 80-40 for females, the Balangodese male 
BP2/17 is stenomeric and the other specimens of the fossil series, both male and 
female, are eurymeric for this index. With regard to the Pilastric Index, the male 
BP4/8 comes nearest the Vedda means of io8-8o for males and 106-90 for females. 
There is a discrepancy in the size of the femora of BP4/8, however. The other 
Balangodese have a Pilastric Index which is outside the upper range observable for 
the Veddas. There is close agreement, however, between the two series as regards 
the Index of Robusticity, that of Vedda males averaging 11-69, ^.nd the females 
averaging 10-80. Osteological features of greatest similarity for the femora of the 
two series occur more frequently at the extremities of the bone rather than at the 
mid-shaft region. To the differences of the shaft noted above should be added the 
greater torsion of the femur among Veddas. Both groups have very thick shafts 
and a pronounced development of the pilaster with the lips of the linea aspera 
separated to a marked degree. A short neck, compressed antero-posteriorly, and 
supporting a spherical head, is common to the proximal ends of the femora of both 
series. The greater trochanter is prominent and the trochanteric fossa is deep. 
The lesser trochanter is large and smooth. A third trochanter is absent. The 
crista hypotrochanterica is of variable development in the femora of both series. 
The distal end is marked with a broad intercondylar fossa for both Veddas and 
Balangodese, and a further example of similarity is afforded by the weak development 
of the adductor tubercle. Hill notes the great backward extent of the medial 
condyle and the anterior development of the lateral condyle of the Vedda femur. 
This trait is reminiscent of the orientation of the condyles of the Balangodese femur. 

The tibia is mesocnemic for the majority of Vedda and Balangodese specimens, the 
index for the former series averaging 65-95 for males and 68-70 for females. None 
of the Balangodese tibias are pathologically affected as are many of the Vedda 
specimens with consequent distortion of their conformation. Only BP2/17 exhibits 
anterior-posterior bowing of the shaft to the extent found among the greater number 
of Vedda tibiae. The lateral prism formed by the transverse section of the Vedda 
tibia is not observable in the fossil series. Muscularity is moderate for bones of 
both series. The Balangodese tibia is somewhat more thick and massive. Torsion 
of the shaft is most prominent among the Veddas. 

The fibular fragment of BP4/8 resembles the Vedda form in its straightness and 
triangular transverse section. However its articular extremities are more promin- 



HUMAN SKELETAL MATERIAL FROM CEYLON 193 

ently developed. In muscularity it is more strongly marked, although the Vedda 
fibula also exhibits some hollowing for the peroneal and tibialis posticus muscles. 

The tali of the two series are very similar in size and muscular development. The 
area for the fibular malleolus is extensive. However, the neck of the talus is broader 
among the Balangodese and the forward extension of the superior articular facet that 
Hill notes as a common featiire of the Vedda talus is not apparent among the fossil 
specimens. The calcaneus is longer and narrower for the Veddas, but the pronounced 
curvature of the sustentaculum tali brings it nearer to the form found in the older 
group. The latter have a cuboid bone that approximates that of the Vedda cuboid 
in its squat form. The articular facets are prominent, but the Vedda cuboid shows 
greater convexity of the plantar surface. 

The Skeleton of the Pelvis. Apart from the greater size of the Balangodese sacrum 
this bone falls well within the range of variation for other features observable in 
Vedda sacra. The platyhieric sacrum of BP2/17 can be matched with several Vedda 
specimens although the average for the series lie between the dolichohieric andsub- 
platyhieric categories. Closer similarity is seen in the uniform curvature and 
hyperbasal form of this bone. Four out of the eleven Vedda sacra examined by 
Hill showed an articulation of three sacral vertebrae with the ilium. The other 
Vedda sacra have only one or two segments connecting with their ilia. The various 
sacral aberrations that Hill finds in his Vedda specimens are not present in BP2/17, 
i.e., incomplete union of sacral vertebrae, failure of closure of the neural arches, and 
spina fissa of the terminal segments. 

The pelvic bones of BP2/17 exhibit a combination of infantile and " paedo- 
morphic " traits which are common for Vedda pelves plus " gerontomorphic " 
features that are characteristic of specimens of Australian aboriginal populations. 
Of the traits of the latter sort, BP2/17 shows prominent muscular attachments, a 
thick and massive ilium, and a size that is at the upper limit of the range of variation 
for the Vedda pelvic dimensions. Like the more gracile innominates of the Veddas, 
however, the male BP2/17 shows a sciatic notch with a fairly high angle, pronounced 
torsion between ilium and ischium and a deep hollowing of the gluteal fossa. The 
ischium of BP2/17 is narrow and long, but the Vedda form is broader and shorter. 
The Height-Breadth Index for the Veddas measured by Hill is 80-50 for males and 
76-60 for females, indices considerably lower than that obtained by the present writer 
for the Balangodese. The mesopellic category of the Veddas — 95-60 for males and 
95*20 for females of Osman Hill's series — also contrasts with the higher index for the 
Balangoda pelvis. The neural foramina in both groups are cordiform. 

The Dentition. The mean values for the sizes of molar and premolar teeth of 
Balangodese and Veddas show that the higher values occur in the former series. 
Where the Balangodese means are lower — as with the bucco-lingual diameter of the 
RM- and mesio-distal diameters of the RM-, LPM^ and RLMi^ — the differences are 
very shght. Only in the latter case are the grinding teeth of both sides of the mouth 
smaller in size. The Balangodese molars and premolars are larger bucco-lingually 
than those of the Vedda series, hence their greater Length-Breadth Index. The 
anterior teeth in the two series show a significant difference from this molar-premolar 



194 HUMAN SKELETAL MATERIAL FROM CEYLOX 

size relationship. The upper canines of the Balangodese are smaller than those of 
the Vedda, but the labio-lingual diameter retains its relatively greater size over the 
mesio-distal length, and the indices are not as low as they are in the Vedda ratios. 
The mandibular canines reflect this situation. The mesio-distal length is greater 
for the maxillary and mandibular lateral incisors of the Vedda series, but the labio- 
lingual diameter shows no difference, save for the RI- which has a higher mean value 
among the Balangodese. The Length-Breadth Index remains higher for the fossil 
series, however. Their mean index for the upper central incisors is less than that 
for the Veddas, due to their smaller labio-lingual diameter. The mesio-distal 
diameter is larger for the Rli but the same for the LIl. The mandibular central 
incisors cannot be compared due to the inadequate size of the sample. The Balango- 
dese LIi^ shows the higher mean labio-lingual and mesio-distal values, but the index 
is below that of the Vedda series. 

None of the specimens of the Vedda series exhibits on the occlusal surface of the 
lower molars the rectangular form, such as that found in BP2/21. However, the 
maxillary teeth in both series show a low incidence of occlusal rectangularity — a 
deviation from the common rounded molar form. The triangular premolar form is 
absent in the maxilla of the fossil series but present among the Veddas. The man- 
dibles of both series have triangular premolars, but the rectangular form predominated 
in the Balangoda series, the round form in the Vedda. Canines are only triangular 
in the maxillae of the Veddas but BP2/20-41 has a RLC which is rectangular. This 
same specimen has a RLC of like pattern. This form is not absent in the Vedda 
mandibular canines although the common pattern is triangular. Some Vedda males 
have triangular upper incisors, but the only form found in the fossil series is rectangu- 
lar. This is the case for upper and lower dentitions. 

The Balangodese exhibit a greater variation in cusp pattern than do the Veddas, 
who offer no examples of molars with 5 or 6 cusps in the maxillary dentition. Cases 
of tricuspid molars are absent in the mandibles of both series, and 6 cusps are found 
in this region only among the Balangodese — the LM^ of BP3/27-34. This variation 
in the molar cusp patterns, particularly in the maxilla, is a peculiar feature of the 
Balangodese dentition. It is further exemplified in the 4-cusp pattern of the RLPM- 
of BP2/20-41. The buccal cusp(s) of the premolars is consistently the higher in 
both series. The Veddas offer no cases of Carabelli's cusp nor the presence of other 
accessory cusps, but both are to be found in the Balangoda series. Crenulation of 
the third molar appears to be a common feature of both series, although it is found 
exclusivel}^ in the females of the Vedda group. The strict correlation of cusp number 
and groove pattern which is universal in the latter series does not hold true with the 
Balangodese. Furthermore, there are no cases of Y4 or -I-5 groove patterns in the 
Vedda series, the 4 cusp and 5 cusp patterns relating to the +4 and Y5 groove pat- 
terns, respectively. In both series the maxillary cusp pattern for molars is +4- 

The greater degree of attrition of the molars and premolars of male specimens, as is 
the situation in the Vedda series, is not encountered in the fossil series where the 
degree of wear is more pronounced among females. More of the posterior teeth of 
the Balangodese exhibit pronounced abrasion than do the corresponding teeth of 



HUMANSKELETALMATERI A LFROM CEYLON 195 






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GEOL. I I, 4 l8 



200 HUMAN SKELETAL MATERIAL FROM CEYLON 

the Vedda series. The condition of the canines is similar in both groups, the lower 
canines being the more heavily worn. There are no incisor teeth in the Balangoda 
series whose abrasion might be described as " slight " or " sub-medium " as is the 
case for the Veddas. The condition of moderate to pronounced wear is the norm for 
this fossil series. Only among Vedda females do the lower molars appear more 
heavily worn : among the Balangodese the more severe forms of mandibular molar 
abrasion are represented by the males. The Vedda male specimens show greater 
maxillary molar abrasion, but for the Balangodese this sort of sexual differentiation 
does not exist. Both groups, however, show greater wear of the upper incisors 
among the males. The assymetrical wear pattern observable in the right and left 
sides of the dentitions of certain Vedda specimens is not apparent in the fossil series. 

The occurrence of shovel-shaped incisors, while present only in the dentitions of 
the females of the Balangoda series, is found in both sexes of the Veddas. Further- 
more the development of this feature is considerably more prominent among the 
Veddas. This trait appears in the lower dentition of a single Balangodese female — ■ 
BP2/21. Lingual tubercles are absent from the anterior teeth of males of both series. 
Among females their presence may or may not be coincident with shovel-shaping. 

With respect to dental anomalies the Balangodese and Veddas exhibit some 
striking differences. Caries and evidence of abscess are absent in the former but not 
infrequent in the latter population. The frequency of ante-mortem tooth loss is 
high for Veddas, negligible in the fossil series. Both populations show cases of 
supernumerary teeth. Among the Balangodese these are found in the molar portions 
of both jaws and for the Veddas there is a single case of a supernumerary tooth in 



Table 12 



Comparative Anthropometric Data: ' 


Fhe M 


\XILLARY 


Molar 


Dentition 










Ml 






W 






m 


















j^ 






j^ 






1 




^ 


f 




N 


f 




> 








MD 


BL 


I 


MD 


BL 


I 


MD 


BL 


I 


Balangodese 


BP2/17 




10 


12 


114-2 


10-5 


12 


114-2 


10 


II-5 


115-0 




BP2/20 






10 


II 


IIOO 


8-5 


12 


141 -I 










BP4/8 






10 


II 


IIO-O 


9 


II 


122-2 


9 


II-5 


127-7 




BPi/6 






IO-5 


12 


114-2 


10 


12 


120-0 


9-5 


II-5 


121 -o 




BP2/21 






II 


12 


109 -I 


10 


13 


130-0 


8-5 


13 


151-9 




BP3/27 






9 


II 


122-2 


10 


IO-5 


105 








Veddas 










8-5 


12 


150-0 


10 


10-6 


107-4 


10 


10-9 


109-4 


Mundas 










9-6 


10-2 


106-2 


8-5 


10 -2 


120-5 








Oreans 










9-2 


10-7 


II6-2 


8-7 


10-7 


122-8 








Male 










10 


10 


lOO-O 


8 


10 


125-0 








Australians 










II-4 


12-8 


105-2 


10-9 


I3-I 


I20-I 


10 


12-3 


123-0 


Sampung 










II -2 


12-8 


II4-2 


10-9 


12-8 


II7-4 


10 


12-4 


124-0 


Gua Cha 




Brothwell 


1960 


IO-3 


I2-I 


II7-4 


IO-2 


12-2 


109-6 


9-3 


12-0 


119-0 


Talgai 




Campbell 


1925 


12-6 


I3-I 


103-9 


II-3 


133 


II7-6 








Keilor 




Adam 1943 


II 


13-2 


120-0 


9-9 


13 


131-3 


9-7 


12 


123-7 


Niah 










IO-2 


12-3 


120-5 


IO-7 


13 


I2I-4 


II-5 


12-4 


107-8 



H U M A N S K E L E T A L M A T E R I A L F R O M C E Y L O N 201 

the mandibular incisive region. Suppression of teeth is not a feature in either series. 
Crowding is found only in the third mandibular molars in the prehistoric group. 
Crowding is not observable among the Veddas who show, however, a high incidence 
of maladjustment for other parts of the dentition. 

The first molar is more commonly the larger in the posterior dental rows of both 
series, but the presence of the third molar as the one of greater size is found more 
frequently in the earlier series, particularly in the mandibles. There are no cases 
of the first molar being the larger in the lower jaws of the Balangoda series, although 
the Vedda series affords several such cases. Where the third molars are absent, the 
second molar is larger than the first. 

The overbite type of occlusion, which is common for jaws of the Vedda series, 
appears to have been present also among the Balangodese, although only BP2/25 
and BP3/27-34 afford evidence of this. 

IV DISCUSSION 

From the anthropometric and comparative analyses of the human remains from 
Bellan Bandi Palassa originate certain problems of interpretation which require 
comment. Regarding the question of the phenotypic affinities of the Balangodese, 
Deraniyagala has entertained several views. His earliest opinion, conceived at a 
time when the only prehistoric Ceylonese hominid specimen was the one from the 
site of Batadomba lena, was that the Veddas were a composite race of several diff ■^rent 
population elements from India which had introduced the Neolithic culture to the 
Island. " The tendency of the Veddas to throw back to two physical types differing 
from other races in Ceylon ..." suggested this hypothesis (Anonymous 1941 : 354). 
Later he remarked that "... the supposedly autochthonous Vaddha of Ceylon 
possibly carries some proportion of the blood of Balangoda man ..." (Deraniya- 
gala I943« : 112), but following the discovery of the specimens from the sites of 
Ravan alia and Telulla he wrote, 

" This race may be termed ' Proto-Vaddoid '. The so-called Vaddahs who 
do not differ in culture, religion, and language from the forest villagers can only 
be regarded as hybrids between this extinct autochthonous Stone Age race 
and the more modern metal using ones of Ceylon. Supporting this view is the 
fact that the so-called Vaddahs display a variety of physical types that have 
puzzled the anthropologists who attempted to study them under the impression 
that they were a distinct race " (Deraniyagala 1955a : 40). 

As to the Sinhalese he conjectures, 

" The blood of Homo sapiens balangodensis however exists in almost equal 
intensity in the colonies of the so-called Vaddahs of today, and among the 
Sinhalese so that in most cases it would be impossible to distinguish a so-called 
Vaddah from a Sinhalese if both were clothed alike and placed in the same village 
. . . the only recognizable autochthonous race in Ceylon is the extinct one in a 
lithic culture phase ..." (Deraniyagala 19556 : 301-302). 



202 HUMANS KELETALM A TERIALFROMCEYLON 

Since the appearance of the fossils from Bellan Bandi Palassa, he has insisted that 
the Balangodese show a combination of Neanderthaloid and Australoid traits that 
he calls " Proto-Australoid ", plus a Negroid element added at a later time (Deraniya- 
gala 1957a : 3, 4). He suggests that the Balangodese replaced the people of the 
" Ratnapura " culture who were the true autochthones of Ceylon and that the 
Balangodese continued to practice their distinctive culture until the fifth century 
A.D. The Veddas are, he concludes, " a degenerate mixed population " of Balango- 
dese and Sinhalese racial elements (Deraniyagala 1958a : 258, igGoa : 96, 108). 

The only other published discussion of the racial affinities of the Balangodese 
known to the present writer is Coon's (1962 : 424-425). This author agrees with 
Deraniyagala that there is a Negroid element in the series, but believes that the 
dominant strain is " Caucasoid " rather than " Australoid " ; although he hastens 
to add that the presence of some " Australoid " features should not be surprising 
considering the location of the Island of Ceylon just south of a " Caucasoid-Australoid 
zone of contact ". He appears to base this opinion upon the presence of reduced 
frontal brow ridges, a sharply pointed chin, the lack of a prominent nuchal crest 
and what he regards as a narrow and prominent nasal structure which are represented 
in the single specimen now on loan at the American Museum of Natural History in 
New York (Specimen T-24-B or BP3/i5b). Although Dr. Coon saw six of the 
Balangodese specimens in Ceylon in 1957, these had not yet been cleaned, repaired 
or described. While the present writer, basing his analysis upon the restored 
specimens, agrees that Balangoda Man did not differ subspecifically from the living 
Veddas, as Coon states, the complete series supports an interpretation that must 
take into consideration the non-Caucasoid phenotypic traits of the Balangodese. 
Since Coon regards the living Veddas as " Caucasoids " along with their Sinhalese 
neighbours, it is not surprising that he has emphasized the racial criteria of this 
phenotype in his interpretation of the Balangodese population. 

Emerging from this very general comparative anah'sis of the Balangoda phenotype 
with the phenotypes of other populations and individual specimens, both prehistoric 
and contemporary, are two major considerations : (i) the high frequency of similar 
morphological traits shared by the Balangodese and the Veddas which suggests a 
positive genetic affinity between them ; (2) the number of unique morphological 
traits that characterize the Balangodese which are among those traits recognized 
by various students of the Veddas as the phenotypic hallmarks of half-breeds or 
" Vedda sub-types ". Hill (1941 : 134) lists the main effects of miscegenation in the 
Vedda population as the tendency to increase of the cranial capacity above 1,300 cc, 
an increase in the relative height and/or breadth of the cranial vault with a consequent 
lessening of the steepness of the lateral walls and the formation of reduced temporal 
fossae, an elongation of the face, especially in the increased depth of the mandible, 
an absence of prognathism, a constriction of the orbits and rounding off of the outlet 
of the orbital fossae, a narrowing of the spheno-maxillary fissure, a reduction in the 
extent of the frontal bone on the inner wall of the orbit, and the presence of leptorrhiny 
which is accompanied by longer nasal bones, an elevated nasion and an oxycraspedote 
apertura pyriformis. In the majority of cases Hill can confirm the mixed 



HUMAN SKELETAL MATERIAL FROM CEYLON 203 

ancestry of certain Vedda crania by facts relating to their history. Indeed the 
proportion of these atypical crania is greatest for those collected since 1886. How- 
ever, Hill is cautious in assigning to these crania on the basis of his observations 
alone anj^ traits that would identify the non- Vedda phenotype per se, i.e., absolute 
traits indicative of admixture from Sinhalese, Tamils, Malays, etc. Nor does he 
regard all crania well outside the modal values for the Vedda phenotype as specimens 
which are non-Vedda in pedigree. Rather, he notes the presence of a third type of 
cranium which is less typical than his " classic Vedda " type but which he cannot 
place with the " half-breed Veddas ". He suggests that these individuals may either 
be the result of remote miscegenation followed by a later addition of Vedda genes 
from unadulterated gene pools, or else representatives from within the range of 
variation of pure Veddas. The skulls of the second type, he notes, exhibit a number 
of traits described as " primitive, even simian " (Hill 1941 : 124, 125, 134). 

This is the most recent of several schemes to account for physical subtypes among 
the Veddas. That of von Eickstedt (1927^) allows for five types : (i) the " Veddoid " 
which corresponds to the conventional " pure Vedda " type living the old way of 
life ; (2) the " Singhalesoid " or acculturated Vedda living in villages ; (3) the 
" Mongoloid " type which shows admixture with Malays ; (4) the " pseudo-Austra- 
loid " whose criterion is a heavier facial beard than that of the Veddoid type ; (5) 
the Coast Vedda who has become mixed with Tamils. Two or three subtypes were 
mentioned by Topinard (1885) after an examination of twelve Vedda crania at the 
Royal College of Surgeons. Hocart (1931 : 5) suggests that there is an element in 
Ceylon's population that has not as yet been identified, and Raghaven (1953 : 51), 
seeing Mediterranean and Australoid elements at opposite ends of the range of 
variation of Vedda morphology, finds a third type. This he claims is shorter in 
stature than the Australoid and Mediterranean and has been called Negritoid. 

This observation that certain physical features found in the Balangodese are 
among those features that anthropologists have held to be the criteria of Vedda 
crosses or sub-types does not weaken the claim that considerable miscegenation of 
the Veddas with the Sinhalese, Tamil and other ethnic groups of Ceylon has trans- 
pired : rather, it attempts to explain how those physical features which cannot be 
attributed to the modern non-Vedda populations happen to be manifested in certain 
proportions among the Veddas of the present day. That the presence of relatively 
compressed temporal fossae and the absence of total facial prognathism in Vedda 
populations are related to their recent genetic crossings cannot be doubted. But 
other features — leptoprosopy, hypsicrany, greater thickness of the cranial bones, 
prominent mastoid development, the peculiar morphology of the mandible — are 
examples of physical features shared by both the Balangodese and the Veddas. 
On the basis of the Balangoda specimen T-24-B the presence of leptorrhiny in the 
Balangodese gene pool cannot be excluded, although specimen BP4/8 indicates that 
platyrrhiny was also a feature of the group. In addition to these traits which have 
been selected as evidence of Vedda miscegenation or penetrance of archaic sub-types 
are a number of others which the Veddas share with the Balangodese but which have 
a very low frequency among any of the Ceylonese populations today. The most 



204 HUMAN SKELETAL MATERIAL FROM CEYLON 

Striking of these are the similarities in cranial conformation, prominence of supra- 
orbital tori, the depth of the nasal notch, platyrrhiny, chaemaeconchy and certain 
similarities of the post-cranial skeleton. Of those physical traits that distinguish 
Balangodese from Veddas as well as from other Ceylonese, the majority appear in the 
post-cranial skeleton. Apart from their greater lengths, the long bones are more 
robust and lack the bowing and torsion that characterizes those of the Veddas. 
Some cranial distinctions are reflected in their greater palate dimensions, pronounced 
mandibular muscularity and a dentition in which the molars are larger in size and 
exhibit cusp patterns of a more complex development. These would appear to be 
physical characteristics that were never manifested in the Vedda phenotype. But 
while the Balangodese may share a similar genetic background with the Veddas, 
they are " pre- Vedda " primarily in the chronological sense of that term. Genetically 
both Balangodese and Veddas appear to have been recipients of a common gene 
pool in the past. Affinities with other phenotypic groups, living and prehistoric, 
cannot be questioned, as the comparative Tables are able to illustrate to a certain 
degree. But of all the populations with which the Balangodese were compared by 
the present describer, none appi cached them in number and significance of mor- 
phological similarities to the degree represented by the Veddas of the historic period 
of Ceylon. 

Historically the Veddas have been treated as though they were a homogeneous 
racial or sub-racial entity which the physical anthropologist could clearly distinguish 
from other ethnic groups living in Ceylon. However a little investigation into the 
problem of how the Veddas have been biologically and culturally defined indicates 
that this subject has been the focus of considerable debate, as attested by the fact 
that certain writers have declared the Veddas to be extinct while their contemporaries 
have estimated that they number in the thousands. It is suggested here that the 
matter of how the concept of the Veddas as a distinctive phenotypic pattern within 
Ceylon came to be is deserving of some careful research in the light of the new osteo- 
logical evidence from Bellan Bandi Palassa. Conceivably such a programme of 
research could be approached along three avenues of investigation : (i) the problem 
of how the term " Veddas " has been employed and the determination of what this 
means when applied to the population as a whole as well as to individuals within the 
population so defined, (2) the problem of how the Veddas have been described in 
scientific and popular literature and in oral traditions, (3) the analj'sis of the various 
interpretations that have been made by those writers concerned with the Veddas 
when applying their data to the wider fields of human evolution and cultural be- 
haviour. 

Regarding the first problem, it must be understood that the concept of the Veddas 
as a relict population with ancient indigenous roots in Ceylon has been a basic 
assumption behind all definitions of their culture and phenotype. For the Sinhalese, 
Tamils, Malays, Chinese, Moormen, Europeans and other ethnic groups of the Island 
there are historical accounts of their colonization, but for the Veddas (and Balango- 
dese) there is no historical documentation of their longer habitation here. Their 
past must be reconstructed from prehistoric archaeology, native Sinhalese and Tamil 



HUMAN SKELETAL MATERIAL FROM CEYLON 205 

chronicles, and anthropometric data. The problem is further complicated by the 
fact that the term Vedda has become defined not only as a distinct physical type 
but on the basis of sociological criteria as well. By the close of the igth Century 
three concepts regarding the physical morphology of the Veddas had been put forward 
(i) the Veddas constituted a single homogeneous racial population morphologically 
distinct from the macro-population of Ceylon ; (2) the Veddas were racially hetero- 
geneous save for " true ", i e., " pure Veddas ", who followed a primitive hunting- 
gathering economy ; (3) the " true Veddas ", whatever their economic status, were 
definable in terms of one or more osteological specimens proclaimed as typical of 
the Vedda phenotype. An additional concept has been offered by Hill (1945 : 
202-203) who considers the geographical area rather than the degree of acculturation 
to be the vital factor in the recognition of a physical type definable as Vedda. 

In scientific and popular descriptions of the Vedda phenotype, selection has been 
the major stumbling block of physical anthropologists. The number of osteological 
specimens collected from Ceylon since 1827 and bearing the name " Vedda " on their 
labels is impressive and exceeds in size the series accessible for many other Asiatic 
tribal groups of higher population frequency, and the amount of anthropometric data 
pertaining to both osseous and living Vedda specimens is not inconsiderable. Never- 
theless this abundance of data has not deceived the more perceptive students of the 
Veddas who have troubled themselves to investigate the histories of particular 
so-called Vedda specimens and Vedda communities. The history of the scientific 
investigation of Ceylon's aboriginal population has yet to be written, but a cursory 
glance at the published data indicates that evidence gleaned from small samples often 
has been considered applicable to the Vedda population as a whole. In most cases 
the fact that the specimens were from Vedda populations at all is questionable, for 
the majority of collectors obtained them through Sinhalese and Tamil contacts by 
offering to pay the latter for every skull they could procure, leaving it up to the con- 
tacts to decide what was or was not a Vedda. The care taken by the Sarasins 
and Hill to get their osseous specimens from Vedda cave sites and cemeteries, 
personally excavated, is a commendable but infrequent condition in the history of 
Vedda osteological collections. Furthermore, even those specimens which most 
experts would recognize as Veddas were taken from a fairly circumscribed sector 
of the eastern portion of the Island, although historical accounts make it clear that 
in the past the Veddas were more widespread if not ubiquitous. 

Scientific research on the Veddas, which may be said to have commenced in the 
early decades of the last century, has never been completely independent of the 
notions contained in fantastic traveller's accounts or of the oral and written traditions 
concerning this population. Even in the rigour of the determination to select those 
Vedda specimens, osseous or living, which are "t3^pical", scientific workers have 
repeatedly emphasized the importance of certain phenotypic traits and thus intro- 
duced additional selective factors into the interpretation of What is a Vedda? Of 
these notions of the Vedda phenotype, the most common is that it is infantile and/or 
anthropoidal and hence representative of a very primitive and even atavistic kind 
of humanity. Some writers have barely accorded the Veddas human status. The 



2o6 HUMAN SKELETAL MATERIAL FROM CEYLON 

selection of certain Vedda crania as prototypes, even though it was an attempt to 
define with clarity the concept of the Vedda phenotype, has tended to obscure the 
factor of normal variability of this population. The ghost of this tradition lingers 
in the works of those researchers who, even in the manipulation of large series of 
Vedda specimens, have not been emancipated from the employment of such terms 
as " typical Veddas ", " pure Veddas " or " classic Veddas " in their writings. Such 
expressions may be useful when denoting specimens for which no admixture with 
non- Vedda phenotypes is suspected, but they are misleading when the total Vedda 
phenotype is the subject of consideration. Such terms are meaningless when the 
physical anthropologist attempts to understand the phylogeny of the Veddas since 
the gene pool of any population is never static. In Ceylon population shifts have 
been especially encouraged through the operation of frequent genetic intercourse 
between groups as well as by the dynamics of genetic drift and natural selection in 
local areas of the Island. 

An examination of the archaeological complex at Bellan Bandi Palassa forces 
the anthropologist to ask these questions : (i) How are the cultural artifacts found 
in association with the Balangoda skeletal series related to the prehistoric picture 
of Ceylon and the Indian mainland and to southern Asia as a whole? (2) What 
affinities, if any, can be demonstrated between the cultures of the Balangodese and 
the historic Veddas? These problems of the archaeological significance of the cxiltural 
assemblage at Bellan Bandi Palassa are complementary to the problems of the physi- 
cal anthropology of the human remains. In comparing certain elements of the 
Balangoda culture with that of the Veddas, attention is directed to a paper by 
Allchin (1959). In summarizing the evidence of the prehistoric cultures of Ceylon, 
AUchin has pointed out striking similarities between certain of its elements and par- 
ticular cultural traits of the Veddas. The present writer, although restricting himself 
to the cultural evidence from the single site of Bellan Bandi Palassa, finds grounds 
for justifying a similar comparison. To the early prehistorians who pioneered 
research into Ceylon's prehistory the suggestion that the ancestors of the Veddas were 
the manufacturers of the ubiquitous stone tools seemed obvious. Those who 
questioned this apparent relationship between the prehistoric or contemporary cul- 
tures did so because they thought the Veddas incapable of producing a lithic industry, 
and not because they questioned the validity of the archaeological evidence in the 
Vedda caves. Now that the physical remains of the artisans of these lithic industries 
have been recovered, the contentions of the Sarasins and the Seligmanns must be re- 
evaluated along with the concepts of their opponents about the role that the ancestors 
of the Veddas played in Ceylon's prehistory. 

As a final point in this discussion it must be mentioned that what is now required 
of the prehistorians of Ceylonese studies is a compilation of data concerning the 
Veddas from ancient native chronicles and later travellers' accounts. Whatever 
the affinities of the Veddas to the prehistoric and historic populations of Ceylon 
might be, the invasion of the Sinhalese in the Fifth Century B.C. caused a concentra- 
tion of the indigenes in the eastern wilderness of the Island — the Veddarata — where 
they are still to be found in isolated chena villages. That they once occupied the 



HUMAN SKELETAL MATERIAL FROM CEYLON 207 

entire Island, as has been suggested by the Sarasins (Sarasin & Sarasin 1907 : i8g, 
F. Sarasin 1926 : 83) cannot be proved as yet, but references to their presence in 
other portions of Ceylon can be inferred from a cursory glance at the native Sinhalese 
and later European records as well as from a mapping of the Vedda place names 
across Ceylon. These data establish that the territory of the Veddas extended far 
beyond the present limits of this relict group within the recent historic period. 
What their area of occupation may have been during the period that Ceylon was 
known only along its coastline and before its interior had been penetrated by Sin- 
halese and Tamil colonization awaits further research. The migration of Vedda 
communities over the Island during the past three millenia and earlier is relevant 
to the problems of their physical morphology today. One wonders what selective 
factors, be these biological or cultural, influenced the preservation of certain Vedda 
populations in the relict areas of the Island while other Vedda groups became in- 
corporated in communities of invading peoples or failed to pass on their genes at all. 
Useful as Hill's concept of Vedda geographical-biological areas may be to the 
physical anthropologist desirous of obtaining " typical Vedda " specimens, it must 
be borne in mind that these individuals are a segment of a much larger Vedda 
gene pool which existed in the past and the factors conducive to their survival are 
unknown. Furthermore we are ignorant of the genetic shifts that have taken place 
within these population segments since the commencement of their isolation. Such 
considerations are relevant to the problem of identifying the Balangodese when one 
considers that this stone-using prehistoric population was in existence at the dawn 
of the period of Sinhalese settlement and that the genetic complex that makes it 
distinctive as a phenotypic entity at this period must have had considerable influence 
upon the genetic composition of the present-day peoples of Ceylon. 

V CONCLUSIONS 

The Balangoda skeletal series from Bellan Bandi Palassa constitutes a unique 
phenotypic pattern present in Ceylon at the dawn of the historic period, a time within 
the second and third millenia B.P. While possessing certain biological features that 
distinguish them from other fossil hominids found thus far, the Balangodese fall 
within the range of polytypic variation representative of post-Pleistocene Homo 
sapiens of the Indian-southeast Asian- Australian gene pool. The phenotypic 
pattern most closely resembling that of the Balangodese is that of the Veddas of 
Ceylon. Similarities are not only quantitatively provocative but striking as well in 
terms of particular isolated physical traits that distinguish both Veddas and Balango- 
dese from other southern Asiatic populations. Indeed, certain of the morphological 
and anthropometric characteristics of the fossil series have been cited by physical 
anthropologists studying the Veddas as indicative of Vedda sub-types or as the 
evidence of their miscegenation with other living racial groups. It would seem 
therefore that both the Balangodese and the Veddas are biologically united through 
their possession in the past of a common gene pool, although it must be stressed 
that the former are " pre- Veddas " in a chronological rather than in any direct 
phylogenetic kind of relationship. Both populations appear to have been subject 



2o8 HUMAN SKELETAL MATERIAL FROM CEYLON 

to separate evolutionary pressures for a long period of time, but for reasons which 
are yet unknown the Balangodese phenotypic pattern did not persist into the historic 
period. It is the variety and nature of physical differences between these two 
populations that suggests their bifurcation from a common stem at a time several 
millenia prior to the occupation of Bellan Bandi Palassa. At the dawn of the 
historic period in Ceylon, the Veddas were in all probability distributed throughout 
the Island, save perhaps along the northern coastline, and as a dispersed population 
their phenotype reflected local variations as a consequence of isolation between the 
segments of their population. That the Veddas are simply the hybrid descendants 
of a crossing of Balangodese with Sinhalese or Tamils cannot be supported on the 
basis of the anthropometric and historic data, although the presence of a yet unidenti- 
fied phenotypic element in Ceylon in the prehistoric period must not be entirely 
excluded from considerations of the Balangodese- Vedda phylogeny. 

The cultural affinities of the Balangodese from Bellan Bandi Palassa are with the 
manufacturers of the lithic industries which have been assigned to the Indian Late 
Stone Age or its Ceylonese manifestation, the Bandarawelian. The number and 
kinds of similarities that exist between the prehistoric and Vedda cultures forcefulty 
suggests the existence of a cultural continuum in Ceylon that extends into the 
historic period. Such traits as the use of iron and Sinhalese weapons by the Veddas 
of recent centuries find their parallels in the acquisition of painted pottery, ground 
stone tools, and perhaps other traits by the hunting-gathering and microlith-using 
Balangodese who may well have been introduced to these by trading peoples from 
coastal Ceylon and the Indian mainland. Further speculations of the biological 
and cultural affinities of this prehistoric population await additional research along 
these lines : (i) a resolution of the problem of what constitutes the Vedda phenotype 
apart from the traditional methods of defining this population, (2) an orientation of 
archaeological investigation in Ceylon that can demonstrate the role that the Vedda's 
have played in the manufacturing of the Island's prehistoric industries, (3) the map- 
ping of Vedda occupation sites throughout the Island with the view of establishing 
the migration patterns and degree of mobility of the Veddas within the period of 
recorded history, (4) the recovery of additional osteological material from Ceylonese 
sites for the purpose of broadening our knowledge of the physical anthropology of 
the region. This present analysis of the Balangodese is an initial attempt to extend 
our understanding of the evolution of human populations during that interim which 
is marked by the close of the Pleistocene and the dawn of the historic period in 
southern Asia. 

VI ACKNOWLEDGEMENTS 

I should like to thank Dr. E. I. White, F.R.S., Keeper of Paleontology, and Dr. 
K. P. Oakley, Deputy Keeper of the Sub-Department of Anthropology, of the British 
Museum (Natural History) for permission to study the collections of human osteologi- 
cal material in their department and for their kind assistance and encouragement. 
Dr. P. E. P. Deraniyagala, Director of the National Museum of Ceylon, Colombo, was 
responsible for the discovery of the Bellan Bandi Palassa skeletal series and a person 



HUMAN SKELETAL MATERIAL FROM CEYLON 209 

to whom I am deeply grateful. I am also greatly indebted to these staff members of 
the British Museum (Natural History) : Mr. A. E. Rixon, Mr. D. R. Brothwell, 
Mr. R. J. Parsons, Mr. G. C. Ross, Mr. M. J. Rowlands, Mrs. EUzabeth Gardiner, 
Mrs. Madeleine Glemser, and Miss Rosemary Powers. My thanks are due to Dr. E. A. 
Mourant of the Lister Institute, London, and to Dr. Virginia Carbonell. Apprecia- 
tion is extended to Professors T. D. McCown, S. L. Washburn and S. F. Cook of the 
University of California at Berkeley ; to Mrs. Audrey Dejournette of Oakland, 
California ; and to mj' wife. Dr. Mary C. Marino. The research programme was 
made possible through a grant from the National Science Foundation, Washington, 
D.C. 

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HUMAN SKELETAL MATERIAL FROM CEYLON 211 

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1955- Analytical methods of dating bones. Advanc. Sci., London, 11:3-8. 



212 HUMAN SKELETAL MATERIAL FROM CEYLON 

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Oslo, 2 : 7-193. 



HUMAN SKELETAL MATERIAL FROM CEYLON 213 

WiCKREMASiNGHE, R. L., Ikin, E. W. & MouRANT, A. E. 1963. Blood gtoups and haemo- 
globins of the Veddahs of Ceylon. /. R. anthrop. Inst., London, 93 : 1 17-124. 

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1951. Prehistory in India : Broadcast Talks on Early Man. ii -f- 39 pp., 16 pis. Poona. 

ZucKERMAN, S. 193©. The Adichanallur skulls. Bull. Madras Govt. Mus., 2 : 1-24, illust. 



PLATE I 

Specimen BP2/17. Unreconstructed calvarium cleared from 
its matrix. Left lateral aspect. 



Bull. B.M. (N.H.) Geol. 11, 4 



PLATE 1 




■mm%. 



5cm 

-I — 1 — 1 — 1 — I — I — I 



PLATE 2 

Specimen BP2/17. Unreconstructed calvarium cleared from 
its matrix. Left lateral aspect. 



Bull. B.M. (N.H.) Geol. 11, 4 



PLATE 2 




5cm 



PLATE 3 

Specimen BP2/17. Partial reconstruction of calvarium. 
Frontal aspect. 



Bull.B.M. {N.H.) Geol. 11, 4 



PLATE 3 




PLATE 4 

Specimen BP2/17. Partial reconstruction of calvarium. 
Left lateral aspect. 



Bull. B.M. [N.H.) Geol. 11, 4 



PLATE 4 





5 cm 



T 1 1 



PLATE 5 

Specimen BP2/17. Partial reconstruction of calvarium. 
Right lateral aspect. 



Bull.B.M. [N.H.) Geol. 11, 4 



PLATE 5 




5 cm 
i""""'i ' I ' ' — r — 'I II 




PLATE 6 

Specimen BP2/17. Partial reconstruction of calvarium. 
Superior aspect. 



Bull. B.M. {N.H.) Geol. 11, 4 



PLATE 6 




PLATE 7 
Specimen BP2/ 1 7. Unreconstructed maxilla. Palatal aspect. 



Bull. B.M. (N.H.) Geol. 11, 4 



PLATE 7 




5 cm 
1 1 I M I I M I I 1 -y 



PLATE 8 
Specimen BP2/i7i. Mandible. Left lateral aspect. 



Bull. B.M. (N.H.) Geol. 11, 4 



PLATE 8 




PLATE 9 
Specimen BP2/i7i. Mandible. Superior aspect. 



Bnll.B.M. (N.H.) Geol. 11,4 



PLATE 9 




PLATE lo 

Specimen BP3/27-34. Partial reconstruction of calvarium. 
Left lateral aspect. 



Bull B.M. (N.H.) Geol. II, 4 



PLATE 10 




PLATE II 

Specimen BP3/27-34. Partial reconstruction of calvarium. 
Right lateral aspect. 



Bull.B.M. [N.H.) Geol. 11, 4 



PLATE II 




PLATE 12 



Specimen BP3/27-34. Partial reconstruction of calvarium. 
Occipital aspect. 



BuU.B.M. (N.H.) Geol. 11, 4 



PLATE 12 




5cm 
|""i"H| 1 1 r 



PLATE 13 

Specimen BP3/27-34. Partial reconstruction of calvarium. 
Superior aspect. 



Btill. B.M. (N.H.) Geol. II, 4 



PLATE 13 




|"" I ' I H | 1 1 1 1 

5cm 



PLATE 14 
Specimen BP3/27-34. Mandible. Superior aspect. 



Bull.B.M. {N.H.) Geol. 11,4 



PLATE 14 




5cm 
iiii|iiii| ( 1 1 I 





PLATE 15 

Specimen BP3/27^34. Radiograph of mandible and the premolar 
and molar teeth. Right lateral aspect. 



Bull.B.M. [N.H.) Geol. 11, 4 



PLATE 15 




PRINTEn IN GREAT BRITAIN 
BY ADLARD Kc SON LIMITED 
BARTHOLOMEW PRESS, DORKING 



h ■ 

0. 



( .: 1 DEC 



SOME NEW BRITISH ALBIAN %^^ 
OSTRACODA 



p. KAYE 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. ii No. 5 

LONDON: 1965 



SOME NEW BRITISH ALBIAN OSTRACODA 




?»4r«^ 



BY 

PETER KAYE, Ph.D. 

Burmah Oil Exploration Co. Ltd., 20 Esplanade, Scarborough 



Pp. 215-253 ; II Plates ; 5 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. ii No. 5 

LONDON: 1965 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 5 of the Geological 
{Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 



Trustees of the British Museum (Natural History) 1965 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued December, 1965 Price £2 10s. 



SOME NEW BRITISH ALBIAN OSTRACODA 

By P. KAYE 



I 

II 
III 



CONTENTS 
Introduction and acknowledgements . . . . . 218 

Location of samples . . . . . . . . .219 

Systematic descriptions . . . . . . . .221 

Suborder Cladocopina . . . . . . . .221 

Family Polycopidae . . . . . . . .221 

Genus Poly cope Sars. . . . . . . . .221 

Polycope nuda sp. nov. ...... 221 

Polycope oweni sp. nov. ...... 222 

Suborder Podocopina. ........ 223 

Family Bairdiidae ......... 223 

Genus Bairdia McCoy . . . . . . . .223 

Bairdia pseudoseptentionalis (Mertens) .... 223 

Genus Pontocyprella Mandelstam ...... 224 

Pontocyprella semiquadrata sp. nov. .... 224 

Family Cypridinae . . . . . . . . .225 

Genus Argilloecia Sars. ....... 225 

Argilloecia valvula sp. nov. ...... 225 

Genus Paracypris Sars. . . . . . . .226 

Paracypris wrothamensis sp. nov. .... 226 

Genus Krausella Ulrich ....... 227 

Krausella sp. ........ 227 

Family Cytherideidae ........ 228 

Genus Clithrocytheridea Stephenson ..... 228 

Clithrocytheridea heslertonensis Kaye .... 228 

Genus Schnleridea Swartz & Swain ..... 228 

Schuleridea dimorphica sp. nov. ..... 228 

Genus Habrocythere Triebel ....... 229 

Habrocythere fragilis Triebel ..... 229 

Genus Dolocytheridea Triebel ...... 230 

Dolocytheridea typica sp. nov. ..... 230 

Family Cytheruridae ........ 231 

Genus Eucytherura Muller ....... 231 

Eucytherura aff. nuda Kaye ..... 231 

Genus Hemicytherura Elofson . . . . . .231 

Hemicythenira euglyphea sp. nov. ..... 231 

Genus Cytheropteron Sars. ....... 232 

Cytheropteron (Cytheropteron) argiita sp. nov. . . . 232 

Cytheropteron [Cytheropteron) milbournei sp. nov. . . 233 

Cytheropteron [Cytheropteron) nanissimum nanissimum 
Damotte & Grosdidier ...... 234 

Cytheropteron {Cytheropteron) nanissimum fenestrata subsp. 
nov. ......... 234 

Cytheropteron [Cytheropteron) lamplughi nom. nov. . 235 

Subgenus Eocytheropteron Alexander ..... 235 

Cytheropteron [Eocytheropteron) protonsa sp. nov. . 235 
Subgenus Infracytheropteron Kaye ..... 236 

Cytheropteron [Infracytheropteron) obscura sp. nov. . 236 



Geol. 11,5 



2l8 



SOME NEW BRITISH ALBIAN OSTRACODA 



IV 



Genus Orlhonotacytheve Alexander . 

Orthonotacythere fordensis sp. nov. . 
Orthonotacyihere ntinutissima sp. nov. 
Orthonotacythere spinifera sp. nov. 
Family Brachycythcridae .... 

Genus Alatacy there Murray & Hussy 

Alatacythere robusta robusta (Jones & Hinde) 

Alatacvthere robusta langi subsp. nov. . 

Family Bythocytheridae ..... 

Genus Monoceratina Roth. .... 

Monoceratina longispina (Bosquet) 

Monoceratina sp. .... 

Family Progoncytheridae .... 

Genus Acrocythere Neale .... 

Acrocythere striata sp. nov. . 
Genus Neocythere Mertens .... 
Subgenus Physocythere Kaye 

Neocythere [Physocythere) tenuis sp. nov. 
Family Protocytheridae ..... 
Genus Veenia Butler & Jones 
Veenia compressa Kaye 
Veenia florentinensis Damotte 
Family Trachyleberididae .... 

Genus Cythereis Jones ..... 
Cythereis angulatoides nom. nov. . 
Cythereis gatyensis Damotte & Grosdidier 
Cythereis glabrella Triebel . 
Cythereis pinhayensis sp. nov. 
Suborder Myodocopina 
Family Conchoeciidae 
Genus Conchoecia Dana 
Conchoecia sp.A 
Conchoecia sp.B 
References 



237 
237 
239 
239 
240 
240 
240 
241 
242 
242 
242 
244 
244 
244 
244 
245 
245 
245 
246 
246 
246 
246 

247 
247 

247 
248 
248 
248 
250 
250 
250 
250 
250 
251 



SYNOPSIS 

Twenty new species and subspecies of Ostracoda are described from the Gault Clay (Middle 
and Upper Albian) of England. Additiona information is given on nine already described species 
and four ostracod synonyms [Clithrocytheridea ventricola Damotte & Grosdidier, Cytheropteron 
punctata Kaye, Cythereis angulata Kaye and Cythereis lamplughi Kaye) are corrected. Four 
forms are left under open nomenclature. 



I INTRODUCTION AND ACKNOWLEDGEMENTS 

A quantitative study of the distribution of Ostracoda in the Enghsh Gault (Middle 
and Upper Albian) has been carried out by the collection of accurately localized 
samples at small vertical intervals in as many Gault exposures as are currently 
available. Over three hundred and fifty samples have been collected, a large propor- 
tion of which have already been analysed working on a 1,500 gram basic starting 
weight of sediment. The project as a whole is only partially complete but a certain 
number of undescribcd species have already been found. The description of these new 
species together with other relevant taxonomic information is the subject of the 



SOME NEW BRITISH ALBIAN OSTRACODA 219 

present paper. Twenty species and subspecies belonging to fifteen genera are des- 
cribed for the first time. Additional information is given on nine already known 
species and four Cretaceous ostracod synonyms are corrected. Four forms are left 
under open nomenclature. The known range of each described species is given and 
its value as an index fossil is noted where possible. A complete list of the Gault 
exposures sampled during the course of the overall project is given. A palaeonto- 
logical zonation of the Middle and Upper Albian based on ammonites is given as 
Text-fig. I. All specimens described and illustrated in this paper are deposited in 
the collections of the British Museum of Natural History (B.M.N.H.) but considerable 
numbers of comparative forms are retained in the author's private collections. 

This study has been carried out during the tenure of a D.S.I.R. Research Fellow- 
ship at the Sedimentology Research Laboratory, Dept. of Geology, Reading Univer- 
sity, and I am extremely grateful to Professor P. Allen for all his help and encourage- 
ment. I would also like to thank many of my colleagues at Reading, particularly 
Dr. A. W. Medd, Mr. G. H. Scott and Mr. D. B. WilHams for help in field work and 
Mr. J. L. Watkins for the photography. I would also like to express my gratitude to 
Mr. H. G. Owen and Mr. R. A. Milbourne whose help concerning Gault stratigraphical 
problems has been of immense value. 

II LOCATION OF SAMPLES 

A list of the localities from which Gault Ostracoda have been obtained is as follows : 
(i) Lower and Upper Gault at the British Portland Cement Go's pit. Small Dole near 

Henfield, Sussex. Grid. Ref. TQ. 218131 

(ii) Lower Gault at the Honey Lane Brickworks, Selbourne, Hampshire. Grid. 

Ref. SU. 768342 

(iii) Lower Gault at the Greatness Lane Brick pit, Sevenoaks, Kent. Grid. Ref. 

TQ. 536578. 

(iv) Lower and Upper Gault at Ford Place pit, Wrotham, Kent. Grid. Ref. 

TQ. 636591 

(v) Upper Gault at the Rugby Portland Cement Go's pit, Paddlesworth, near 

Maidstone, Kent. Grid. Ref. TQ. 695623 

(vi) Upper Gault at Sandown Bay and also Blackgang, Isle of Wight 

(vii) Upper Gault at Pinhay Point, Devon. Grid. Ref. SV. 342928 

(viii) Lower Gault at Devizes, Wiltshire. Grid. Ref. ST. 986612 

(ix) Lower Gault at Culham, near Abingdon, Oxfordshire. Grid. Ref. SV. 510949 

(x) Lower and Upper Gault at Mundays Hill pit, Leighton Buzzard, Bedfordshire. 

Grid. Ref. TL. 915978 

(xi) Upper Gault at the London Brick Go's pit, Arlesey, Bedfordshire. Grid. Ref. 

TL. 185352 

(xii) Upper Gault at Fisons Brickpit, Burwell, Cambridgeshire. Grid. Ref. TL. 

516691 

(xiii) Upper Gault at Eastwoods Cement pit, Barrington, Cambridgeshire. Grid. 

Ref. TL. 394507 



SOME NEW BRITISH ALBIAN OSTRACODA 



ALBIAN ZONES and SUB ZONES 



ca 

_j 
< 

DC 

a. 

0. 

3 



Stoliczkaio dispor 



Morloniceras inflatum 



Mortoniceros permflatum 
Arrophoceros substuderi 

Mortoniceros inflatum 
vor. aequotorlolis 

Collihoplites Quritus 

Hysteroceros vorlcosum 

Hy steroceros orbignyi 



< 

_l 
< 

LlI 

_I 



Euhoplites loutus 



Hoplites dentotus 



/ Dipoloceros cristotum 
Anohoplites doviesi 
Euhoplites nitidus 
Euhoplites meondrinus 
Dipoloceros subdeloruei 
Dimorphoplites doris 

Dimorphoplites niobe 
Anohoplites intermedius 
Hoplites spathi 
Hoplites benettignus 
Hoplites eodentotus 



< 

m 

_j 
< 

oc 

UJ 



Douvilleiceros mommillotum 



Leymeriello tordefurcoto 



Protohoplites puzosionus 
Otohoplites roulinionus 
Cleoniceros floridum 
Sonnerotio kitchini 

Leymeriello reguloris 
Hypoconthoplites milletioides 
Fornhomia fornhomensis 



SOME NEW BRITISH ALBIAN OSTRACODA 221 

(xiv) Lower Gault at Castles Farm pit, near Ely, Cambridgeshire. Grid. Ref. 

TL. 600775 

(xv) Lower Gault at Speeton, E. Yorkshire. Grid. Ref. TA. 150758 

(xvi) Lower Gault at West Heslerton, E. Yorkshire. Grid. Ref. TA. 913759 

(xvii) Lower and Upper Gault at Folkestone, Kent. Grid. Ref. TR. 242365 

(xviii) Upper Gault at Ashford, Kent. Grid. Ref. TR. 058435 

(xix) Gault at Swanage, Lulworth, Osmington and Black Ven near Lyme Regis, 

Dorset. 

Many other pits, particularly in the Lower Gault of the Wealden area have been 

found to be barren of Ostracoda. These include : 

(i) Lower Gault at Hassocks, Sussex. Grid. Ref. TO. 310155 

(ii) Lower Gault at the Marley Tile pit, Storrington, Sussex. Grid. Ref. TQ. 

094138 

(iii) Lower Gault at Nyewood Brick pit, Nyewood, near Petersfield, Hampshire. 

Grid. Ref. SU. 800218 

(iv) Lower Gatdt at Wrecclesham, near Farnham, Hampshire. Grid. Ref. SU. 

826448 

(v) Lower Gault at Arnold's sand pit, Buckland, Kent. Grid. Ref. TQ. 231512 
(vi) Lower Gault at Squerry's pit, Westerham, Kent. Grid. Ref. TQ. 442538 
(vii) Lower Gault at Ufhngham, Oxfordshire. Grid. Ref. SU. 315905 
(viii) Lower Gault at Badbury Wick, near Swindon, Wiltshire. Grid. Ref. 

SU. 192818 

(ix) Lower Gault at Thame, Oxfordshire. Grid. Ref. SP. 691055 

(x) Lower Gault at Coney Hill Sandpit, Oxted, Surrey. Grid. Ref. TQ. 375526 

III SYSTEMATIC DESCRIPTIONS 

Suborder CLADOCOPINA 
Family POLYGOPIDAE 

Genus POLYCOPE Sars 1866 
Polycope nuda sp. nov. 
(PI. 4, figs. 1-3) 
Derivation of name, nuda^ — referring to the lack of surface ornament. 

Diagnosis. Large moderately inflated Polycope with subcircular outline and 
smooth valve surface. 

HoLOTYPE : A single left valve, B.M.N.H. lo. 2847, from the Lower Gault niobe 
Subzone ; Wrotham, Kent. 

Paratypes : B.M.N.H. lo. 2848-2850. Three carapaces from the same sub- 
zone and locality. 



222 some new british albian ostracoda 

Measurements. 

Length Height Width 
(mm.) (mm.) (mm.) 

Left valve (B.M.N. H. lo. 2847, holotype) . 0-55 0-50 o-i6 
Carapace (B.M.N.H. lo. 2848) . . .0-58 0-50 0-32 

Description. Valves large, moderately inflated, subcircular in outline. There 
is a slight flattening dorsally along the hinge line forming two weak cardinal angles. 
The anterior margin is more bluntly rounded than the posterior margin. Lateral 
surface mainly smooth but with some weak irregular reticulation particularly antero- 
ventrally in certain specimens. Overlap strong particularly posteriorly and ventrally. 
Hinge simple with a groove in the margin of the right valve to accommodate the 
sharp edge of the left valve. Duplicature not seen. Muscle scars, a triangular 




Fig. 2. Muscle scars of Polycope nuda sp. nov. X2500. 

patch composed of three scars, the apex pointing dorsally. The two ventral scars 
are roughly triangular in shape, the dorsal scar is diamond shaped, fitting between 
the points of the other two scars. 

Remarks. Only one other species of the genus Polycope has been previously 
recorded from the Cretaceous. This species, P. bonnemai Herrig 1964, is known from 
the Maastrichtian of the Isle of Rugen and a similar form Polycope sp. was described 
by Bonnema (1940) from the Maastrichtian of the Netherlands. P. nuda is therefore 
the first pre-chalk Cretaceous reference to the genus and is closely related and 
presumably ancestral to P. bonnemai. It is closely similar in size and ornamentation 
to the latter but differs in details of the shape and overlap, lacking the anterior and 
postero-dorsal marginal thickening of P. bonnemai. In shape the anterior and pos- 
terior margins are more evenly rounded in P. mida. P. oweni described below, also 
from the British Albian, is much smaller and strongly ornamented. The genus is 
fairly common in the Jurassic where several species are known, particularly from the 
Liassic and Oxfordian. In general these forms are smaller, much more flattened and 
usually strongly ornamented. P. nuda is found consistently throughout the Gault 
first making its appearance in the niobe Subzone. It is never very common though its 
relative abundance in the niobe Subzone is a useful indicator of that horizon. 

Polycope oweni sp. nov. 
(PL 4, figs. 11-15) 

Derivation of name. After H. G. Owen whose work on Gault stratigraphy has 
been invaluable to me in my studies of the distribution of the Ostracoda in the Gault. 

Holotype. A left valve, B.M.N.H. lo. 2859, from the H. orbignyi Subzone 
(Upper Gault) ; Wrotham, Kent. 



SOME NEW BRITISH ALBIAN OSTRACODA 223 

Paratypes. B.M.N.H. Io. 2858, 2860-63. Four valves and two carapaces from 
the same horizon and locaUty. 

Diagnosis. Small subcircular Polycope with valve reticulately ornamented. 
Along and at the junction of the reticulation are numerous small spines. 

Measurements. 

Length Height Width 
(mm.) (mm.) (mm.) 

Left valve (B.M.N.H. Io. 2859, holotype) . 0-42 0-37 o-ii 
Carapace (B.M.N.H. Io. 2858) . . .0-42 0-37 0-22 

Description. Valves small, weakly inflated, subcircular in outline. The valves 
have a slightly greater length than height but there is only very slight flattening 
along the hinge line. The valves are slightly asymmetrical anteriorly but the 
anterior and posterior margins are without cardinal angles. Marginal protuberances 
are absent but the whole of the margin is thickened. The lateral surfaces are 
strongly but variably ornamented ranging from reticulation to spination. The 
basic ornament is the reticulation covering the whole of the surface on a somewhat 
concentric pattern. Small spines are developed along the ridges of the reticulations 
giving rise to a completely spined appearance in many of the specimens. Overlap 
is marked all round the margin except dorsally. 

The hinge is simple ; the margins in both valves being flattened to form sharp 
marginal bars which overlap each other. Dorsally in the right valves there is a weak 
groove above the bar. Muscle scars, three equal-sized oval scars arranged in a 
triangle. One apex of the triangle points dorsally. 

Remarks. P. oweni is not abundant in the Gault and is found rarely in clays of 
post A. intermedius Subzone age. It differs from P. nuda in the details of the shape, 
hinge, muscle scars and ornamentation and is smaller. 

Suborder PODOCOPINA 

Family BAIRDIIDAE 

Genus BAIRDIA McCoy 1844 

Bairdia pseudoseptentrionalis (Mertens) 

(PI. 2, figs. I, 3-6) 

? 1840 Cytherina subdeltoidea Munster 



Roemer : 105, pi. 15, fig. 22. 

Reuss : 16, pi. 5, fig. 38. 

Jones : 23, pi. 5, figs. 15a-/. 

Reuss : 140, pi. 26, figs. ^a-c. 

Jones & Hinde : 5, pi. 2, figs. 31-34. 

Alexander, pi. 6, figs. 2, 4. 

Alexander : 61, pi. 3, fig. 5. 
1956 Bairdoppilata pseudoseptentrionalis Mertens : 182, pi. 8, figs. 7-10, pi. 13, figs. 89-90. 
1956 Bairdoppilata roemeri Deroo : 1509, pi. i, figs. 9-12. 
1958 Bairdoppilata pseudoseptentrionalis Mertens ; Howe & Laurencich : 82. 
1958 Bairdoppilata ? roemeri Deroo ; Howe & Laurencich : 82. 



1845 Cytherina subdeltoidea Munster 

1849 Bairdia subdeltoidea (Munster) 

1874 Bairdia subdeltoidea (Munster) 

1890 Bairdia subdeltoidea (Munster) 

1927 Bairdia subdeltoidea (Munster) 

1929 Bairdia subdeltoidea (Munster) 



224 SOME NEW BRITISH ALBIAN OSTRACODA 

Material. B.M.N.H. Io. 2828-2832, 4 valves and i carapace from the Gault 
just below the Cambridge Greensand at Arlesey, Beds. 

Measurements. 

Length Height 
(mm.) (mim.) 

Left valve (B.M.N.H. Io. 2828) . . . . 1-32 0-87 

Right valve (B.M.N.H. Io. 2829) . . . . 1-34 0-75 

Remarks. For a long time all Cretaceous species of Bairdia were placed in 
B. siibdeltoidea (Munster) a Tertiary form. In 1956 Deroo re-named the Cretaceous 
references to B. subdeltoidea as Bairdoppilata roemeri. Earlier in the same year 
Mertens had, however, erected a new species Bairdoppilata pseudoseptenfrionalis 
which seems to be identical with Deroo's form. Both authors had referred their 
species to the genus Bairdoppilata which is said to differ from true Bairdia in having 
two rows of denticles at either end of the hinge line in the right valves. However, 
as van Morkhoven (1963) has pointed out, the presence or absence of such denticles is 
a characteristic of most of the genera within the Bairdiidae each genus containing 
species both with and without them. It therefore seems unwise to perpetuate the 
genus Bairdoppilata. 

Bairdia pseudoseptentrionalis occurs rarely in the Lower Gault but is more abun- 
dant in the Upper Gault. 

Genus PONTOCYPRELLA Mandelstam 1956 
Pontocyprella semiquadrata sp. nov. 
(PI. 3, figs. 1-8) 
Derivation of name. Semiquadrate-alluding to the shape. 

Diagnosis. Pontocyprella with semiquadrate shell. Anterior margin semi- 
circular, posterior margin bluntly rounded. 

HoLOTYPE. A left valve, B.M.N.H. Io. 2834, from the H. orbignyi Subzone, 
Upper Gault ; Wrotham, Kent. 

Paratypes. B.M.N.H. Io. 2835-2837. Three valves from the same horizon and 
locality. 

Measurements. 

Length Height 
(mm.) (mm.) 

Left valve (B.M.N.H. Io. 2834, holotype) . . 0-70 0-40 

Right valve (B.M.N.H. Io. 2836) . . . . 0-73 0-40 

Description. Valves elongate, compressed, semiquadrate in shape. Dorsal 
margin weakly convex, without cardinal angles ; ventral margin weakly concave. 
Anterior margin semicircular, posterior margin bluntly rounded forming a weak 
bulge at f height. Greatest height and width at mid-length. Lateral surface 
smooth. Inner lamella broad with large anterior and posterior vestibules. DupU- 



SOME NEW BRITISH ALBIAN OSTRACODA 225 




Fig. 3. Muscle scars of Pontocyprella semiquadrata sp. nov. X2500. 

cature narrow, crossed by numerous short straight fine radial pore canals. Normal 
pore canals few, small and irregularly scattered. Muscle scars a central rosette of 4 
scars. Hinge simple. In the left valve there is a marginal bar with a long narrow 
shelf-like furrow below it. In the right valve there is a narrow marginal shelf with a 
high bar below it. 

Remarks. Pontocyprella semiquadrata occurs rarely in the upper part of the Lower 
Gault and more commonly in the Upper Gault. Its overall shape and particularly 
the shape of the anterior and posterior margins distinguish it from other Cretaceous 
species of the genus P. superba Neale 1962 and P. rara Kaye 1965 have an acute 
posterior end whilst P. harrisiana Jones is larger, more elongate and has the anterior 
margin bulged dorsally and the posterior margin bulged ventrally. P. semiquadrata 
is closest to P. mandelstami Kaye 1965 differing in that the latter is kidney shaped 
rather than semiquadrate. P. maynci Oertli 1958 has the dorsal margin strongly 
arched. 

Family CYPRIDIDAE 

Genus ARGILLOECIA Sars 1866 

Argilloecia valvule sp. nov. 

(PI. 7, figs. 20-25) 

Derivation of name, valvulus L. = husk. 

Diagnosis. Argilloecia with arched dorsal margin and acute posterior margin. 

HoLOTYPE. A right valve, B.M.N.H. lo. 2914, from the H. orbignyi Subzone, 
Upper Gault ; Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2915-2920. Four valves and two carapaces from the 
same horizon and locality. 

Measurements. 

Length Height 
(mm.) (mm.) 

Right valve (B.M.N.H. lo. 2914, holotype) . . 0-52 0-20 

Left valve (B.M.N.H. lo. 2915) . . . . 0-48 o-i6 

Description. Valves, thin, small, elongate, laterally compressed. Dorsal 
margin strongly arched, without cardinal angles ; ventral margin long and straight. 
Anterior margin forming a blunt point antero-ventrally ; posterior margin meeting 
ventral margin at an angle of 90° ventrally. Greatest height just in front of mid- 
length ; greatest width just behind mid-length. Right valve larger than left, over- 



226 SOME NEW BRITISH ALBIAN OSTRACODA 

lapping around the entire margin but particularly strongly antero-dorsally and 
postero-dorsally. Lateral surface smooth. Duplicature narrow, crossed by few 
short, straight radial pore canals. Inner margin and line of concrescence separate, 
forming large vestibules anteriorly and posteriorly. Normal pore canals few, ir- 
regularly scattered. Hinge simple consisting of a groove in the dorsal margin of the 
right valves to accommodate the dorsal edge of the smaller left valves. 
Muscle scars a central rosette of five scars. 



Fig. 4. Muscle scars of Argilloecia valvula sp. nov. X2500. 

Remarks. Argilloecia valvula occurs rarely in the Upper Gault and the upper part 
of the Lower Gault. It is the first record of the genus in the Lower Cretaceous. The 
larger right valve separating the genus from most Cyprids, a feature only otherwise 
occurring in Macrocypris and Pontocypris. Argilloecia differs from these latter two 
genera in size, hinge and marginal features. The large size of the known Albian 
species of Macrocypris easily differentiates them from A. valvtila. M. parva Kaye 
1965 from the Hauterivian and Barremian of Northern England is similar in size but 
differs appreciably in shape. 



Genus PARACYPRIS Sars 1866 

Paracypris wrothamensis sp. nov. 

(PI. 9 figs. 9-14) 

Derivation of name. After the village of Wrotham, Kent the type locality for 
the species. 

Diagnosis. Large Paracypris with strongly drawn out acute posterior end and 
high strongly angular anterior cardinal angle. 

Holotype. a left valve B.M.N.H. lo. 2959 from the H. orhignyi Subzone 
(Upper Albian) ; Wrotham Kent. 

Paratypes. B.M.N.H. lo. 2955-58, 2960, 2961. Three left valves, three right 
valves and two carapaces from the same horizon and locality. 

Measurements. 



Left valve (B.M.N.H. lo. 2959, holotype) 

Right valve (B.M.N.H. lo. 2958) .... 

Carapace (B.M.N.H. lo. 2957) . . . . 0-90 0-35 



Length 


Height 


(mm.) 


(mm.) 


0-92 


0-35 


0-90 


0-35 



SOME NEW BRITISH ALBIAN OSTRACODA 227 

Description. Valves large, compressed, very elongate. Greatest height at I 
length, greatest width at ^ length. In lateral view elongate subtriangular with the 
dorsal margin straight and the ventral margin weakly concave. Anterior margin 
broadly rounded forming a marked cardinal angle with the strongly sloping dorsal 
margin. Posterior drawn out forming a strongly acute point postero-ventrally. A 
well marked postero-dorsal cardinal angle occurs. Lateral surface smooth. Left 
valve larger than right overlapping particularly dorsally and ventrally. 

Large crescentic anterior and large triangular posterior vestibules occur. Zone of 
fusion very narrow anteriorly and posteriorly, doubling in width ventrally. Crossed 
by few thick short radial pore canals branching distally (8-10 anteriorly). Normal 
pore canals small, few, irregularly scattered. Hinge simple, the dorsal edge of the 
right valve fitting into a narrow shelf-like groove along the dorsal margin of the left 
valve. An anterior prolongation of the dorsal marginal bar of the left valve forms a 
tooth-like extension corresponding with a slight incurving of the margin in front of the 
anterior cardinal angle in the right valve. Muscle scars a central rosette of five or 
six oval scars. 

Remarks. P. wrothamensis occurs throughout the Gault in England. It first 
appears in the intermedius Subzone but is rare in the Lower Gault. It becomes 
abundant in the Upper Gault and its abundance is a useful indicator of Upper Gault 
age. It is strongly related to the Lower Cretaceous species P. acuta (Cornuel) 1848. 
This latter species is poorly described and rather confused but is smaller and not so 
acute posteriorly as P. wrothamensis. P. sinuata Neale 1963 from the Hauterivian 
and Barremian is probably, at least in part, synonymous with P. acuta. P. sinuata 
differs from P. wrothamensis in being lower, having a less well marked anterior 
cardinal angle and being much less acute. Of other Cretaceous species P. depressa 
Bonnema 1940 and P. jonesi Bonnema 1940 have the cardinal angles rounded and the 
greatest height further forward ; P. gracilis (Bosquet) and P. siliqua Jones & Hinde 
1890 differ markedly in outline. 



Genus KRAUSELLA Ulrich 1894 
Krausella sp. 

(PL 3, figs. 15, 16) 

Material. Two right valves B.M.N.H. lo. 2845-2846, from the upper part of 
the Lower Gault at Castles Farm, Ely. 

Measurements. 

Length Height 
(mm.) (mm.) 
Right valve (B.M.N.H. lo. 2845; . . . . 0-55 0-27 

Remarks. Only one other species of this genus {K. minuta Triehel) is known 
from the Cretaceous. It is smaller and is not so pointed posteriorly. The strong 
inequality of the valves distinguishes the genus. 



228 SOME NEW BRITISH ALBIAN OSTRACODA 

Family CYTHERIDEIDAE 

Genus CLITHROCYTHERIDEA Stephenson 1936 

Clithrocytheridea heslertonensis Kaye 

(PI. I, figs. 8-12) 

1963a Clithrocytheridea heslertonensis Kaye : 30, pi. i, figs. 10-13. 

1963 Clithrocytheridea ? ventricola Damotte & Grosdidier : 53, pi. i, figs. la-f. 

Material. B.M.N.H. Io. 2825-2827, Female left and right valves, and carapace 
from Bed N. 5, Middle Albian spathi Subzone, West Heslerton, East Yorks. 

Remarks. Specimens of Clithrocytheridea ? ventricola kindly sent to me by Dr. E. 
Grosdidier show that this form is synonymous with C. heslertonensis Kaye. Clithro- 
cytheridea heslertonensis is found abundantly in the H. spathi Subzone of the Lower 
Gault and appears to be confined to that subzone. It is a valuable zonal index and 
has been recorded from West Heslerton, Speeton, Devizes, Culham and Henfield in 
clays of that age. 

Genus SCHULERIDEA Swartz & Swain 1946 
Schuleridea dimorphica sp. nov. 
(PL 5, figs. 1-6) 
Derivation of name. Alluding to the strongly dimorphic nature of the species. 

Diagnosis. Small strongly dimorphic Schuleridea with weak ocular sulcus and 
without cardinal angles. 

HoLOTYPE. A male left valve, B.M.N.H. Io. 2864, from the H. orbignyi Subzone, 
Upper Gault ; Wrotham, Kent. 

Paratypes. B.M.N.H. Io. 2865-2870. Twelve valves and two carapaces from 
the same horizon and locality. 

Measurements. 

Length Height 
(mm.) (mm.) 

Male left valve (B.M.N.H. Io. 2864, holotype) . 0-64 0-41 

Female left valve (B.M.N.H. Io. 2867) . . . 0-52 0-40 

Male right valve (B.M.N.H. Io. 2866) . . . 0-63 0-35 

Female right valve (B.M.N.H. Io. 2868) . . 0-50 0-34 

Description. Valves relatively small, ovate, laterally compressed. Dorsal 
margin strongly arched in left valves, weakly arched in right valves. Cardinal angles 
absent from both valves. Ventral margin convex in the left valves, straight in the 
right. Anterior margin broadly rounded ; posterior bluntly pointed just below mid- 
height. Greatest height just in front of mid-length, greatest width at mid-length. 
Eye spot very weak with a short, shallow, oblique sulcus posterior to it. Lateral 
surface smooth. There is a slight ventral tumidity particularly in the right valves. 
Duplicature broad, crossed by numerous fine radial and pseudoradial pore canals. 
These canals are bent upwards anterodorsally. No marginal denticulation in 
either valve. Inner margin and line of concrescence coincide throughout Normal 



SOME NEW BRITISH ALBIAN OSTRACODA 229 

pore canals abundant and irregularly scattered over the lateral surface. Hinge 
strongly developed, having in the left valves two terminal divided sockets, open 
ventrally, separated by a shelf-like furrow. Dorsal of the median furrow is a high, 
smooth bar and a narrow, shallow accommodation groove. In the right valves there 
are two bar-like denticulate terminal teeth (5 denticles in each) separated by a low, 
median bar. Above the hinge is a prominent median shelf. 

Sexual dimorphism is particularly strong. 

Remarks. Schuleridea dimorphica occurs abundantly in the Upper Gault and is a 
valuable index form for that age. Its small size and ovoid shape distinguish it from 
most other species of the genus. It is closest to 5. sulcata Kaye 1965a from the Aptian 
of the Isle of Wight, differing in the poorer development of the eye tubercle and ocular 
sulcus and in the shape of the dorsal and posterior margins. Smaller and differing 
in hingement from the only other known Albian species S. jonesiana (Bosquet) 1852, 
it is also flatter and lacks the cardinal angles and prominent eye tubercle of S. 
virginis Grosdidier 1964 and the truncate posterior end of S. bernouilensis Grosdidier 
1964. In shape and dimorphic features it differs also from S. derooi Damotte & 
Grosdidier 1963a. 

Genus HABROCYTHERE Triebel 1940 
Habrocythere fragilis Triebel 

(PI. 6, figs. 7-13) 

1940 Habrocythere fragilis Triebel : 166, pi. i, figs. 10-13, pl- 9. ^g- loi- 
1956 Habrocythere fragilis Triebel; Mertens : 198, pi. 10, figs. 51-52. 
1963 Habrocythere fragilis Triebel ; Kaye : 33, pi. 3, figs. 8-9. 

Material. Four normal specimens B.M.N.H. lo. 2882, 2884, 2887, 2888, from the 
D. cristatum Subzone (Middle Albian) ; Wrotham, Kent. 

Three anomalous specimens, B.M.N.H. lo. 2883, 2885, 2886, from the same subzone 
and locality. 

Remarks. Habrocythere fragilis is common throughout the English Albian. 
Certain specimens have, however, been found that may throw light upon the origin 
of this monotypic genus. Amongst the normal assemblages of this species certain 
specimens occur which are identical to the species in its general accepted sense but 
have a large eye spot mounted on a large tubercle set antero-dorsally. They also 
have the ventro-lateral portion of the valves drawn out into a weak alate expansion. 
These specimens have to be included in H. fragilis on the basis of their other features 
and their close association with normal specimens of the species but are closely similar 
to species of the genus Eur yitycy there. It is therefore, very likely that Euryitycythere 
known from Valanginian to Aptian strata has evolved into the genus Habrocythere 
with certain specimens of the latter genus showing recapitulation. The major 
diagnostic features of Euryitycy there are the eye tubercle, the inflated alate ventro- 
lateral area, the wide flattened marginal area and the hinge. The wide flattened 
marginal area and associated radial canals are found in all specimens of Habrocythere 
whilst the eye tubercle and inflation are shown in the " recapitulation " specimens. 



230 SOME NEW BRITISH ALBIAN OSTRACODA 

The variability of the hinge in Ostracoda and its modification during phylogeny is 
well known in Ostracoda and modification during the Hauterivian-Albian time interval 
is quite feasible. The Aptian occurrence of the genus (Kaye 1965(2) is of a single closed 
carapace and the hinge features are not known. Slight simplification of the hinge 
structure during the phylogeny can perhaps be related to the small size and light build 
of Habrocythere and can be used as evidence against making Euryiiycythere Oertli 1958 
a synonym of Habrocythere Triebel 1940. 

Specimens of H. fragilis showing affinities with Euryiiycythere have been found 
from several localities. They occur mainly in the daviesi and cristatum Subzones at 
Wrotham and in the subdelaniei Subzone at Sevenoaks. They are useful indicators for 
recognizing the upper part of the Lower Gault and it is possible that ecological con- 
ditions in the Weald at this time stimulated this diversification. 

Genus DOLOCYTHERIDEA Triebel 1938 

Dolocytheridea typica sp. nov. 

(PI. 3, figs. 9-14) 

Derivation of name. Typicus L. = typical. 

Diagnosis. Small Dolocytheridea, inflated posteriorly, with dorsal margin strongly 
arched postero-dorsally. Posterior end rather truncated. 

HoLOTYPE. A left valve, B.M.N.H. lo. 2839, from the basal Upper Gault (Upper 
Albian) ; Pinhay, Devon. 

Paratypes. B.M.N.H. lo. 2838, 2840-44. Eight valves and two carapaces, 
from the same horizon and locality. 
Measurements Length Height 

(mm.) (mm.) 

Left valve (B.M.N.H. lo. 2839 holotype) . . 0-59 0-33 

Right valve (B.M.N.H. lo. 2838) . . . . 0-57 0-28 

Description. Valves small, elongate, laterally inflated particularly posteriorly. 
Dorsal margin strongly arched, without cardinal angles ; ventral margin straight. 
Anterior margin broadly rounded, posterior margin bluntly rounded rather truncate. 
Greatest height and width at f length. Lateral surface smooth, eye spots absent. 
Duplicature broad, crossed by a very large number of thin, rather sinuous, simple 
radial pore canals. Inner margin and line of concrescence coincide throughout. 
Normal pore canals fairly abundant, irregularly scattered. Muscle scars, not usually 
seen, appear to consist of a vertical row of four oval scars with a V-shaped scar anterior 
to them. Other smaller scars probably occur above and below the main group. The 
hinge is simple consisting in the left valves of a curved smooth groove deepened at its 
ends accommodating the dorsal edge of the smaller right valve. The dorsal margin 
of the right valve is enlarged to form weak smooth terminal teeth. 

Remarks. This species is much smaller than most other species of the genus. It 
differs significantly from the even smaller Z). minuta Kaye inshape and greater inflation. 



SOME NEW BRITISH ALBIAN OSTRACODA 231 

From the D. hilseana — D. intermedia — D. bosquetiana lineage it differs in the greater 
inflation posteriorly and the truncation of the posterior end as well as in size. 

D. typica has so far only been found from the basal Upper Gault at Pinhay in 
Devon. 

Family CYTHERURIDAE 

Genus EUCYTHERURA MuUer 1894 

Eucytherura aff. nuda Kaye 

(PL 7, figs. 17, 18) 

Material. B.M.N.H. Io. 2912-13. Two specimens from the H. orbignyi Subzone, 
Upper Gault ; Wrotham, Kent. 

Remarks. Specimens similar to the Barremian form E. mida Kaye 1964 occur 
rarely in the Upper Gault. They are comparable with true E. nuda but have the 
postero- ventral lobe more strongly inflated and a stronger surface reticulation. It is 
almost certain that extra material will show that the Gault specimens belong to a 
separate species or subspecies. A single valve of a related but distinct form has been 
found from the uppermost Lower Gault at Castles Farm, Ely. This specimen is 
similar in shape and size to E. nuda but has two rows of low surface nodes, one row 
dorsally and a more prominent one ventrally. 

Genus HEMICYTHERURA Elofson 1941 
Hemicytherura euglyphea sp. nov. 
(PL 8, figs. 1-4) 
Derivation of name, euglypheus L. = distinctly marked. 

Diagnosis. Sexually dimorphic Hemicytherura with strong surface ornament of 
longitudinal ridges. 

HoLOTYPE. A male left valve, B.M.N.H. Io. 2921, from the H. orbignyi Subzone, 
Upper Gault ; Wrotham, Kent. 

Paratypes. B.M.N.H. Io. 2922-2925. Five valves and one carapace from the 
same horizon and locality. 

Measurements. 

Length Height 
(mm.) (mm.) 

Male left valve (B.M.N.H. Io. 2921, holotype) . 0-40 0-19 

Female left valve (B.M.N.H. Io. 2924) . . . 0-33 o-i6 

Male right valve (B.M.N.H. Io. 2923) . . . 0-40 0-20 

Female right valve (B.M.N.H. Io. 2922) . . 0-33 0-17 

Description. Valves small, elongate, laterally compressed. Dorsal margin well 
arched in the right valves, weakly arched in the left valves ; without cardinal angles. 

Geol. II, 5 23 



232 SOME NEW BRITISH ALBIAN OSTRACODA 

Ventral margin straight. Anterior margin broadly rounded ; posterior drawn out 
into a short acute caudal process just above mid-height. Greatest height and width 
at mid-length. Lateral surface strongly ornamented, tumid ventrally. Eye spot 
distinct, low and glassy. Ornament consists of a series of longitudinal ridges joined 
by weaker vertical cross ridges. The pattern of longitudinal ridges is not regular in 
the centro-lateral area where a flattened irregularly ridged patch occurs. The strong 
ornament covers the whole of the valve exterior, even anteriorly where several 
longitudinal ridges run strongly across the marginal area. 

Duplicature moderately broad, crossed by few straight, simple radial pore canals. 
There is a well developed ocular pit antero-dorsally in the interior of the valves. Hinge 
rather complex. In the right valve there are two faintly denticulate terminal bar- 
like teeth. Above each tooth is a shelf-like furrow accommodating the dorsal edge 
of the left valve. Between and in line with the teeth and furrows the valve remains 
open but above the general line of the hinge there is a median dorsal marginal bar. 
In the left valve the dorsal margin is enlarged to form a curved marginal bar, more 
prominent at the ends where it fits above the terminal teeth of the right valve. Below 
this marginal bar both anteriorly and posteriorly there are shelf-like terminal 
grooves to accommodate the terminal teeth of the right valve. Sexual dimorphism 
well marked. 

Remarks. H. euglyphea occurs rarely in the upper part of the Lower Gault 
and more commonly in the Upper Gault, being distinguished from most other species 
of related genera on account of shape, hinge and ornament. It is closest to Cytherura 
reticulosa (Chapman) which itself is probably a Hemicytherura, differing in being less 
arched dorsally, in lacking the alate spine postero-ventrally and in details of the 
surface ornamentation. H. euglyphea differs from other Cretaceous species such as 
H. unisulcata Veen, H. bisulcata Veen and H. asiculcata Veen in lacking vertical 
sulcation and in surface ornamentation. 

Genus CYTHEROPTERON Sars 1866 
Cytheropteron fC.J arguta sp. nov. 
(PL 8, figs. 12-17) 
Derivation of name, arguta L. = distinct. 

Diagnosis. Cytheropteron with strongly reticulate lateral surface and ridged 
posteriorly pointing alate expansion. 

HoLOTYPE. A left valve, B.M.N.H. lo. 2936, from the H. orhignyi Subzone, 
Upper Gault ; Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2933-2935, 2937-2939. Six valves and one carapace 
from the same horizon and locality. 
Measurements. 

Length Height 

(mm.) (mm.) 

Left valve (B.M.N.H. lo. 2936 holotype) . . 0-42 0-26 

Right valve (B.M.N.H. lo. 2933) . . . . 0-42 0-24 



SOME NEW BRITISH ALBIAN OSTRACODA 233 

Description. Valves small, ovate, strongly ornamented. Dorsal margin 
strongly arched, without cardinal angles ; ventral margin weakly convex. Anterior 
margin bluntly rounded, posterior drawn out into a caudal extension at mid-height. 
Greatest height and width at mid-length. A rounded ventro-lateral alate expansion 
occurs. Along its crest there is a prominent ridge starting anteriorly at the margin 
at J height and terminating posteriorly in a small posteriorly directed spine. The 
anterior end of the ridge forms a spine on the margin. The lateral surface is strongly 
reticulate. The ventral undersurface is weakly striated. Duplicature moderately 
broad, crossed by few straight, thick radial pore canals. Normal pore canals and 
muscle scars not seen. Hinge very strongly developed with two long, almost 
interdentate, terminal sockets separated by a short straight coarsely crenulate bar 
in the left valve. Above the hinge is a wide marginal shelf. In the right valve there 
are two terminal rows of denticles (6 or 7 in each) decreasing in height towards the 
median element and separated by a short interdentate furrow. Above the hinge is a 
narrow marginal shelf. 

Remarks. Cytheropteron (C.) argiita appears to be confined to the Upper Gault. 
It is extremely abundant in the H. orbignyi Subzone and can be used as an index 
species. It has been found so far at Wrotham, Henfield and Pinhay. 

The strength of the ornamentation and hinge make C. (C) arguta distinct from 
other known species as does the shape of the alae. It is closest to C. (C.) piinctata 
Kaye from the Barremian of Northern England which has a similar ridged alae. The 
latter, however, has the ridge separated from the anterior margin and lacks the strong 
ornamentation. 

Cytheropteron fC.J milbournei sp. nov. 
(PI. 7, figs. 4, 6-9) 

Derivation of name. After R. H. Milbourne whose stratigraphical work on the 
Gault has been an invaluable assistance to my study of the distribution of Ostracoda 
in the Gault. 

Diagnosis. Cytheropteron with drawn out postero-ventrally directed alate 
expansion and punctate ornament over whole lateral surface. 

HoLOTYPE. A left valve, B.M.N.H. lo. 2898, from the subdelaruei Subzone, Lower 
Gault ; Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2899, a left valve from the same horizon and locality 
as the holotype ; B.M.N.H. lo. 2900-02. Two valves and one carapace from the 
niobe Subzone at Sevenoaks. 
Measurements. 

Length Height 
(mm.) (mm.) 

Left valve (B.M.N.H. lo. 2898, holotype) . . 0-37 0-22 

Right valve (B.M.N.H. lo. 2900) . . . . 0-35 0-22 

Description. Valves small, elongate, laterally compressed and with a prominent 
ventro-lateral alate expansion. Dorsal margin weakly arched ; ventral margin 



234 SOME NEW BRITISH ALBIAN OSTRACODA 

straight. Anterior margin semicircular, posterior end drawn out into a blunt caudal 
extension at f height. Greatest height in front of mid-length ; greatest width at 
fan J th . Dorsal margin without cardinal angles. Alae directed postero-ventrally 
and bearing a weak vertical sulcus on its crest. Whole of lateral surface covered with 
a series of pits. The ventral underside of the alae bears a few short longitudinal 
riblets. Duplicature moderately broad, crossed by few thick, straight radial pore 
canals (5 anteriorly). Hinge merodont consisting of a long straight denticulate 
median bar and two small divided terminal sockets in the left valve. Above the 
median element is a wide marginal shelf. 

Remarks. Cytheropteron (C.) milbournei is known from the top part of the Lower 
Gault at Sevenoaks and Wrotham and it is never very abundant. It differs from 
C. (C.) nanissimum principally in the shape of the alae and in its ornamentation. 

Cytheropteron (C) nanissimum nanissimum Damotte & Grosdidier 

(PI. 7, figs. 13, 15) 

1963 Cytheropteron (C.) nanissumum Damotte & Grosdidier : 56, pi, i, figs. 2a-f. 

Material. B.M.N.H. Io. 2907-08, two left valves from the H. orbignyi Subzone, 
Upper Gault ; Wrotham, Kent. 

Remarks. This subspecies occurs throughout the Gault, appearing first in the 
niobe Subzone but not becoming abundant until the Upper Gault. 

Cytheropteron f C.J nanissimum fenestrata subsp. nov. 
(PL 7, figs. 14, 16. 19) 

Derivation of name, fenestrata L. = window. 

Diagnosis. Subspecies of Cytheropteron (C.) nanissimum with fenestrate 

ornamentation on ventral alae. 

HoLOTYPE. A left valve, B.M.N.H. 2910, from the H. spathi Subzone, Lower 
Gault ; Henfield, Sussex. 

Paratypes. B.M.N.H. Io. 2909, 2911. Two left valves from the same horizon 
and locality. 

Measurements. 

Length Height 
(mm.) (mm.) 

Left valve (B.M.N.H. Io. 2910, holotype) . . 0-33 0-20 

Description. Valves small, elongate with a ventro-lateral alate expansion. 
Dorsal and ventral margins convex, greatest height at mid-length. Anterior margin 
semicircular, posterior margin bluntly pointed forming a weak cardinal angle 
dorsally. Alae directed, posteriorly, with a small spine at its end and a weak vertical 
sulcus on its upper surface. Above the alae near the dorsal margin is a low node. 
From this node a rib runs downwards to the crest of the alae , whilst a short longitudinal 



SOME NEW BRITISH ALBIAN OSTRACODA 235 

rib runs across the ala at mid-height of the valves. The two ribs cross on the alae 
and form a " window hke " pattern of ornament. The rest of the lateral surface is 
smooth. 

Internal features identical with C. nanissunium s.s. 

Remarks. This subspecies is closely similar to C. (C.) nanissimuni s.s. differing 
only in the nature of the ornamentation on the alae. In C. nanissiimum s.s. the 
dorsal node and vertical rib are much stronger than in C. (C.) nanissumum fenestrata 
whilst the cross rib is absent. 

■ The limited occurrence of C. (C.) nanissumum fenestrata in the H. spathi Subzone 
points to an ancestral relationship to C. nanissumum s.s. which does not appear until 
the^. niobe Subzone. C. (C) reightonensis Kaye 1964 from the Barremian of Northern 
England is also very closely related being ornamented with rows of pits along the alae. 

Cytheropteron (Cytheropteron) lamplughi nom. nov. 

1964 Cytheropteron (C.) punctata Kaye : 103, pi. 5, figs. 7-8. 

Remarks. Professor W. A. Van den Bold of Louisana State University has kindly 
pointed out that the form I erected from the Lower Barremian at Speeton is in 
synonomy with Cytheropteron punctatum Brady (1868 : 449). I have therefore 
renamed my form Cytheropteron (C.) lamplughi in honour of G. W. Lamplugh and his 
basic research on the British Lower Cretaceous. 

Subgenus EOCYTHEROPTERON Alexander 1933 

Cytheropteron (Eocytheropteron) protonsa sp. nov. 

(PL 6, figs. 1-6) 

Derivation of name. Protonsa L. = stretched out. 

Diagnosis. Large Cytheropteron with bilobed wing-like alate expansion and 
prominent upturned posterior caudal process. 

Holotype. a left valve, B.M.N.H. lo. 2879, from the D. cristatum Subzone, 
Lower Gault ; Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2880-81. Two valves from the same horizon and 
locality. 

Measurements. 

Length Height 
(mm.) (m.m.) 

Left valve (B.M.N.H. lo. 2879 holotype) . . 0-70 0-42 

Right valve (B.M.N.H. lo. 2880) . . . . 0-67 0-42 

Description. Valves large, elongate, laterally compressed. Dorsal margin 
strongly arched ; without cardinal angles in the left valves but with weak cardinal 
angles in the right valves. Ventral margin straight totally obscured by the promi- 
nent ventral alate expansion. Anterior margin broadly rounded ; posterior elon- 



236 SOME NEW BRITISH ALBIAN OSTRACODA 

gated into a prominent upturned caudal process at f height. A large wing-like 
bilobed alate expansion occurs ventro-laterally ; the posterior lobe being more 
extended. The lateral surface is covered with irregular vertical riblets which are 
only poorly developed anteriorly but are stronger posteriorly. The ventral under- 
surface bears a series of longitudinal ridges. A weak eye node occurs antero-dorsally. 
Duplicature rather narrow, crossed by few, straight, simple radial pore canals. 
Inner margin and line of concrescence coincide. Normal pore canals few. Hinge 
strongly developed. In the left valve there is a long interdentate furrow widest and 
deepest at the ends. In the right valve there is a long curved row of denticles, 
highest at the anterior and posterior ends. 

Remarks. C. [E.) protonsa occurs rarely in the upper part of the Lower Gault 
and in the Upper Gault. The wing-like bilobed alae, strong upturned caudal pro- 
cess, hinge and ornament make it quite unlike any other described species of Cytherop- 
teron s.l. 

Subgenus INFRACYTHEROPTERON Kaye 1964 
Cytheropteron ? finfracytheropteronj obscura sp. nov. 
(PL 7, figs. 10-12) 

Derivation of name, obscura L. = obscure. 

Diagnosis. Ovate Cytheropteron s.l. with smooth lateral surface and rounded 
ventro-lateral tumidity. Hinge simple. 

HoLOTYPE. A left valve, B.M.N.H. lo. 2903, from the H. orbignyi Subzone, Upper 
Gault, Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2904-06. Three valves and two carapaces from the 
same locality and horizon. 

Measurements. 

Length Height 

(mm.) (mm.) 

Left valve (B.M.N.H. lo. 2903 holotype) . . 0-42 0-25 

Right valve (B.M.N.H. lo. 2905) . . . . 0-40 0-25 

Description. Valves small, smooth, ovate with a marked ventro-lateral tumidity. 
Dorsal margin strongly arched, without cardinal angles. Ventral margin short and 
straight. Anterior margin broadly rounded, posterior drawn out into a blunt 
caudal process at just over mid-height. Greatest height at \ length, greatest width 
at mid-length. Lateral surface strongly and evenly inflated but with a flat marginal 
shelf anteriorly, posteriorly and postero-ventrally. A low, smooth eye tubercle 
occurs antero-dorsally. Duplicature moderately broad, particularly anteriorly and 
postero-ventrally, crossed by few straight, thick radial pore canals (7-8 anteriorly). 
Inner margin and line of concrescence coincide. Normal pore canals small and few. 
Hinge simple consisting of a smooth marginal bar in the left valve which fits on a 
marginal shelf in the right valve. Below the marginal shelf is a narrow median bar 
being discontinuous anteriorly and posteriorly. 



SOME NEW BRITISH ALBIAN OSTRACODA 237 

Remarks. C. (/.) obscura occurs throughout the bulk of the Gault. It first 
appears in the niobe Subzone but only becomes common in the Upper Gault. The 
relationship of the species is rather problematical, and it has been placed in the genus 
Cytheropteron s.l. on account of its shape and size. The lack of a marked alate 
expansion makes it akin to the subgenus Eocytheropteron but the hinge places it in 
the subgenus Infracytheropteron. The species shows strong similarities to the 
Tertiary genera Bythocythere and Loxoconcha. It differs primarily in the hingement 
and lack of vestibules though it does have a wide marginal shelf anteriorly and 
postero-ventrally. It is closest to Loxoconcha minuta Jennings 1936 from the 
Maastrichtian of New Jersey which is similar in shape and size but has ornamented 
valves. This latter form is doubtfully referred to the genus Loxoconcha. 

Genus O RT HON OTACY THERE Alexander 1934 

Orthonotacythere fordensis sp. nov. 

(PL 5, figs. 7-13) 

Derivation of name. After Ford Place near Wrotham, Kent, the location of the 
holotype. 

Diagnosis. Orthonotacythere with weak but distinct ribbing, low tubercles and 
wide deep median sulcus. The postero-lateral area is reticulate and strongly inflated. 

Holotype. A male left valve, B.M.N.H. lo. 2871, from the intermedius Subzone, 
Lower Gault, Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2872-78. Six valves and two carapaces from the 
same horizon and locality. 

Measurements. 



Male left valve (B.M.N.H. lo. 2871, holotype) 
Male right valve (B.M.N.H. lo. 2872) . 
Female left valve (B.M.N.H. lo. 2873) . 

Description. Valves elongate, compressed, divided by 
median sulcus. Dorsal margin straight, ventral margin convex. Anterior broadly 
rounded, posterior forming a blunt postero-dorsal process. The median sulcus is 
limited ventrally by an irregular longitudinal ridge, below which, on the ventral 
undersurface, lies a further straight longitidunal ridge. Faint cross riblets join the 
two ribs. Antero-dorsally there is a prominent glassy eye tubercle jointed by a short 
rib to a reticulate tubercle immediately below it. Two ridges run from this lower 
tubercle, one anteriorly to meet the anterior margin, the other posteriorly towards the 
sulcus to meet a small tubercle where it turns abruptly through 90° to run vertically 
to connect with an irregular tubercle at the anterior end of the upper ventral longitu- 
dinal ridge. The area between the ridges is reticulate and irregularly ribbed. The lower 
ventral ridge runs anteriorly to reach the margin at \ height and posteriorly to fade 



Length 


Height 


(mm.) 


(mm.) 


0-57 


0-30 


0-55 


0-28 


0-50 


0-30 


, wide 


deep vertical 



238 SOME NEW BRITISH ALBIAN OSTRACODA 

out on the postero- ventral surface. The upper ventral ridge bears two weak tubercles 
ventrolaterally, the posterior one being more prominent. Faint irregular ridges run ver- 
tically from these tubercles merging with the general reticulation of the postero- 
lateral surface. This reticulation is made up of a series of weak longitudinal ridges 
joined by cross members. Antero-dorsally on the postero-lateral surface lies a low 
ridged tubercle. Internal features and hinge typical of the genus. 




Fig. 5 . Diagrammatic representation of the distribution of the ornamentation of Orthonotacythere 

fordensis sp. nov. x 75 

Remarks. 0. fordensis occurs fairly commonly in the A. intermedius Subzone of 
the Lower Gault. It seems to be restricted to that subzone and is a valuable index 
form. . fordensis is closely related to 0. inversa (Cornuel) and related species form- 
ing an evolutionary sequence in the British Lower Cretaceous. It has all the major 
features of the group differing in minor details of the ornamentation and particularly 
in the subdued nature of the costation and tuberculation. Though almost certainly 
derived from these Lower Cretaceous forms it does not, however, show continuation 
of the trends seen there. The tendency towards increased tuberculation and sim- 
plification of costation is not followed and the species must therefore not be on the 
main "Boreal" stock. The related species of Orthonotacythere found in the "Tethyan" 
province do not follow the same trends during evolution, and ornamental patterns 
anomalous to that seen at Speeton have already been found (Kaye 1965(2) in the Aptian 
of the Isle of Wight. It is to these species that close ancestry of 0. fordensis must be 
attributed. The ornamentation of these Aptian species such as 0. catalaunica 
Damotte & Grosdidier and 0. atypica Kaye is closely similar to 0. fordensis having the 
costation fairly complex with particularly the irregular nature of the upper ventral 
longitudinal ridge and the nature of the antero-dorsal costation in good agreement. 
The absence of 0. fordensis s.s. from the Lower Gault at Speeton and its replacement 
by fragmentary specimens of a form intermediate between 0. fordensis and 0. inversa 
tuber culata seem to bear this out particularly on consideration of the joining of the 
" Boreal " and " Tethyan " seas in this area in Apto-Albian times. 

Only a single specimen belonging to this group of the genus Orthonotacythere has so 
far been found from post H. spathi age Gault deposits. This specimen from the 
nitidus Subzone (Bed 31) at Henfield shows increased tuberctilation, poor costation, 
poor development of the ventral longitudinal ridge and no surface reticulation. 
These features seem consistent with the overall trends seen at Speeton and the 
specimen perhaps shows a further stage of this major evolutionary pattern. 



SOME NEW BRITISH ALBIAN OSTRACODA 239 

Orthonotacythere minutissima sp. nov. 

(PL 8, figs. 5-11) 

Derivation of name, minutissimum L. = very small. 

Diagnosis. Very small Orthonotacythere with ventral alate longitudinal rib and 
prominent eye tubercle. 

HoLOTYPE. A left valve, B.M.N.H. lo. 2926, from the H. varicosum Subzone, 
Upper Gault, Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2927-32. Five valves and one carapace from the same 
horizon and locality. 
Measurements. Length Height 

(mm.) (mm.) 
Left valve (B.M.N.H. lo. 2926, holotype) . . 0-28 o-i6 

Right valve (B.M.N.H. lo. 2927) . . . . 0-28 0-15 

Description. Valves very small, elongate, compressed. Dorsal margin long and 
straight ; ventral margin straight, shorter and parallel to it. Anterior margin 
bluntly rounded ; posterior with a weak caudal process at the postero-dorsal cardinal 
angle. Eye tubercle exceedingly prominent, protruding well above the dorsal 
margin. Greatest height at i length at the anterior cardinal angle ; greatest width 
just behind mid-length. Lateral surface divided by a vertical median sulcus which is 
limited ventrally by a prominent alate longitudinal ridge. This ridge increases in 
height posteriorly, being drawn out into a blunt laterally directed spine. The 
base of the sulcus does, however, notch this rib slightly, weakly dividing it into two 
sections. A large tubercle occurs dorsally on the postero-lateral area. Lateral 
surface smooth or pitted. Anterior margin denticulate (5 denticles). Duplicature 
moderately broad, crossed by few, straight radial pore canals. Inner margin and 
line of concrescence coincide. Interior eye pit prominent. Hinge simple consisting 
of a long, straight denticulate bar in the left valve. In the right valve there is a long, 
straight interdentate furrow with a high smooth bar above it. 

Remarks. 0. minutissima occurs rarely in the Upper Gault but has not yet been 
found in the Lower Gault. Its exceptionally small size makes it distinct from other 
species of the genus. The pattern of the main ornamentation shows no affinities to 
any other described form but I have found identical or closely comparable forms from 
the Hauterivian/Barremian of Lincolnshire. The hinge of this species is not strictly 
typical of the genus Orthonotacythere, lacking well defined terminal elements, but the 
shape, sulcus and ornamentation are all closely comparable to that genus. It may, 
however, be possible to include this species in a redefined subgenus such as Stillina 
Laurencich. 

Orthonotacythere spinifera sp. nov. 

(PI. 7, figs. 1-3, 5) 

Derivation of name. Spinifera L. = spined. 

Diagnosis. Small Orthonotacythere with weak median sulcus, pronounced reticula- 
tion over whole of valves, and a series of large spines arranged over lateral surface. 



240 SOME NEW BRITISH ALBIAN OSTRACODA 

HoLOTYPE. A left valve, B.M.N.H. lo. 2895, from the A. intermedius Subzone 
(Bed 13), Lower Gault, Wrotham, Kent. 

Paratypes. B.M.N.H. lo. 2894, a right valve from the H. orbignyi subzone (Bed 
13), Upper Gault, Wrotham. B.M.N.H. lo. 2896-97, two valves from the A. 
intermedius Subzone, Lower Gault, Henfield, Sussex. 

Measurements. Length Height 

(mm.) (mm.) 

Left valve (B.M.N.H. lo. 2895, holotype) . . 0-42 0-22 

Right valve (B.M.N.H. lo. 2894) . . . . 0-42 0-22 

Description. Valves small, elongate, strongly laterally compressed. Dorsal 
margin long and straight ; ventral margin short, straight and parallel to it. Anterior 
margin bluntly rounded forming a well marked antero-dorsal cardinal angle ; pos- 
terior drawn out to form an acute postero-dorsal caudal process. Anterior and 
posterior margins strongly spined. Vertical median sulcus very weak or absent. 
Eye tubercle rounded and glassy, set at dorsal end of a short high, blade-like verti- 
cally elongated spine. A similar bladed spine lies at | length on the dorsal margin 
with a further smaller spine at ^ length just below the dorsal margin. An arcuate 
row of four extremely prominent spines runs parallel to the ventral margin. Below 
this row is a series of smaller semi-fused spines culminating posteriorly in a larger 
bilobed spine. A pair of smaller spines occur posteriorly, one on the dorsal margin, 
the other being vertically below it at f height. The whole of the lateral surface is 
strongly reticulate. 

Duplicature narrow, crossed by few straight radial pore canals. Normal pore 
canals and muscle scars not seen. Hinge strongly developed consisting in the left 
valve of two short, subdivided sockets separated by a long, straight, lobed bar. 
Above the median bar is a narrow marginal shelf. In the right valve there are two 
bar-like denticulate terminal teeth (5 denticles in each) separated by a wide, strongly 
divided, median groove. 

Remarks. Orthonotacy there spinifera is rare in most of the Gault, a few specimens 
having been found at a variety of levels and localities. It is more abundant in the 
Gault at Pinhay in Devon and is perhaps a littoral species. 0. spinifera is quite 
unlike other described species of the genus having such characteristic features as the 
weak median sulcus entirely covered by the surface reticulations and the series of 
marginal and lateral spines. It is smaller than most other species of the genus but 
does show some affinities with the 0. inversa group in the arrangement of the spines. 

Family BRACHYCYTHERIDAE 

Genus ALATACYTHERE Murray & Hussy 1942 
Alatacythere robusta robusta (Jones & Hinde) 
(PL 10, figs. 9, 10) 
19646 Alatacythere robusta (Jones & Hinde) ; Kaye : 51, pi. 2, fig. 18. 

Material. B.M.N.H. lo. 2964, 2965, two valves from the uppermost Gault 
immediately below the Cambridge Greensand, Arlesey Beds. 



some new british albian ostracoda 
Measurements. 



Length 
(mm.) 



Left valve (B.M.N.H. lo. 2965) 



241 



Height 
(mm.) 



1-05 0-51 



1-02 


0-54 


0-98 


0-53 


0-92 


0-50 


0-95 


0-50 



Alatacythere robusta langi subsp. nov. 

(PI. 10, figs. 1-4) 

Derivation of name. After W. D. Lang in appreciation of his stratigraphical 
and palaeontological work on the Cretaceous in Devon and Dorset. 

Diagnosis. Subspecies of Alatacythere robusta (Jones & Hinde) 1890 having 
ventro-lateral alae less drawn out and dorsal longitudinal ridge more pronounced. 

Holotype. a male left valve, B.M.N.H. lo. 2940, from the basal Upper Gault ; 
Pinhay, Devon. 

Paratypes. B.M.N.H. lo. 2941-44. Six valves from the same horizon and 
locality. 

Measurements. 

Length Height 
(mm.) (mm.) 

Male left valve (B.M.N.H. lo. 2940, holotype) . 
Male right valve (B.M.N.H. lo. 2943) 
Female Right valve (B.M.N.H. lo. 294:^; . 
Female left valve (B.M.N.H. lo. 2941) . 

Description. Valves large, elongate, laterally compressed. Dorsal and ventral 
margins straight, converging slightly posteriorly. There are well marked antero- 
dorsal and postero-dorsal cardinal angles. Anterior margin broadly rounded ; 
posterior margin weakly pointed at mid-height, both strongly denticulate. Each 
marginal tubercle corresponds to the extremity of a radial pore canal. Greatest 
height at I length ; greatest width at J length. Eye tubercle large and glassy, 
joined to a prominent anterior marginal ridge which follows the entire anterior 
margin to be continued along the crest of the high extended ventral alate expansion. 
This alate expansion obscures the whole of the ventral margin and strongly increases 
in height posteriorly where it terminates in a short ventro-laterally directed spine. 
The ventral undersurface lacks marked costation. A short, high longitudinal ridge 
occurs obscuring the posterior and central part of the dorsal margin. It is not con- 
tinued to join the eye tubercle. The lateral surface is smooth. Duplicature moder- 
ately broad, crossed by numerous straight radial and pseudoradial pore canals. 
Inner margin and line of concrescence coincide throughout. Normal pore canals 
scarce over the bulk of the lateral surface but abundant along the ridged crest of the 
alae. Hinge strong, amphidont, consisting in the left valve of two terminal sockets, 
open ventrally and separated by a long, straight, denticulate bar. Below the 
anterior end of the median bar there is a high, smooth, circular tooth, whilst above the 



242 SOME NEW BRITISH ALBIAN OSTRACODA 

whole of it there is a narrow marginal shelf but no accommodation groove. In the 
right valve there is a high " boss-like " anterior terminal tooth and a triangular, 
elongate, divided posterior tooth separated by a long straight locellate furrow. 
Anteriorly the median groove opens into a deep smooth circular socket. Above and 
below the median groove are narrow bars. 

Remarks. A robiista langi has so far been found from the basal Upper Gault, 
H. orhignyi Subzone at Pinhay, Devon, at Swanage, Dorset and in the Isle of Wight. 
It occurs earlier than A. robusta s.s. which is found in the topmost Gault of the 
Wealden Area and East Anglia. A . robusta s.s. is distinguished from A . robusta langi 
by the greater lateral elongation of the alae and the weak development of the dorsal 
ridge which does not obscure the margin. The greater elongation of the alae makes 
the valves higher. A. robusta langi is the earliest occurrence of the genus in the 
Cretaceous and specimens of Cythereis reticulata s.l. Jones & Hinde from the top 
Lower Gault have been found at Henfield showing weak ventral alation and typical 
Alatacythere shape though retaining the reticulation. Such forms are possibly 
intermediates between the two genera and indicate the origin of the genus Alatacythere. 

Family BYTHOCYTHERIDAE 

Genus MONOCERATINA Roth 1928 
Monoceratina longispina (Bosquet) 
(PI. I, figs. 3-7) 

1 854 Cyihere longispina Bosquet: 86. pi . 6. figs ja d 

1941 Monoceratina longispina (Bosquet); Bonnima: 40, pi. 6 fihs 67-76 

1964& Monoceratina longispina (Bosquet); Kaye: 53, pi. 3 fig i 

1964 Monoceratina longispina (Bosquet); Szczechura: 388, pi, 3, fig 5 pi. 11, fig. i: 

Material. B.M.N.H. Io. 2820-23 from the H. orbignyi Subzone, Upper Gault ; 
Wrotham, Kent. B.M.N.H. Io. 2824 from the E. doris Subzone, Lower Gault ; 
Wrotham, Kent. 

Measurements. 



Adult left valve (B.M.N.H. Io. 2820) 

Adult right valve (B.M.N.H. Io. 2821) . 

Juvenile carapace (B.M.N.H. Io. 2822) . 

Juvenile left valve (B.M.N.H. Io. 2823) . 

Description. The finding of additional material allows a fuller description of this 
species than that given by Kaye (19646). 

Valves elongate, inflated laterally. Dorsal and ventral margins straight and 
parallel. Cardinal angles well developed. Anterior margin broadly rounded ; 
posterior triangular, angled postero-dorsally. Lateral surface inflated with a 
prominent vertical median sulcus. Below the sulcus is a broad based conical, 



Length 


Height 


(mm.) 


(mm.) 


0-82 


0-42 


o-8o 


0-38 


0-65 


0-35 


o-'?6 


0-^2 



SOME NEW BRITISH ALBIAN OSTRACODA 243 

laterally directed spine. This spine and associated swelling limit the sulcus strongly. 
Apart from a few rows of faint concentric reticulation along the crest of the alae the 
lateral surface is smooth. Some faint reticulation does, however, occur on the 
swollen antero-lateral part of the valves in certain specimens. The greatest width is 
just posterior to the mid-point of the valves. The marginal parts of the valves are 
rather flattened particularly anteriorly and ventrally. 

The hinge consists of a long narrow groove in the right valve which accommodates 
the long, straight, smooth marginal bar of the left valve. Weak false sockets are 
sometimes developed, particularly anteriorly in the left valve. The porcellaneous 
preservation of the specimens obtained make the radial pore canals and muscle scars 
invisible, normal pore canals are, howevei, rather small and irregularly scattered over 
the lateral surface. 

Juveniles are rather dissimilar to the adults. In general they have a rounded alate 
expansion and no spine. This difference is, however, seen in the adults of certain 
species of Monoceratina (particularly from the British Oxfordian) and is possibly a 
dimorphic feature. By no means all the juveniles of M. longispina are just alate and a 
few specimens with spines have been found. The occurrence of specimens without 
spines is not common to all species of Monoceratina and no specimens without 
spines belonging to M. umbonata Williamson have been found in the Gault even though 
the species is much more abundant than M. longispina. The juveniles of M. longis- 
pina are themselves variable in ornamentation and two instars are illustrated. Penul- 
timate instars have a wide marginal flattened area. The inflated alate expansion is 
strongly reticulate and a large node covered with reticulation occurs anterior to the 
sulcus. A narrow ridge crosses the sulcus and runs on to the postero-lateral area. 
Smaller instars are much less inflated and lack the flattened marginal area. They 
are completely smooth and have the sulcus wider and more open. Besides being 
strongly limited v^entrally the sulcus is weakly limited dorsally giving the form of a 
wide median depression. 

Remarks. This species is not common but occurs consistently throughout the 
Gault. The original description was from the Senonian and I have recorded it from 
the Cambridge Greensand at Barrington. It has so far been found in the Gault of the 
Wealden area at Folkestone, Wrotham, Sevenoaks and Henfield where specimens occur 
in clays ranging in age from A. intermedhis Subzone to H. varicosum Subzone. It 
has not been found in the spathi Subzone assemblages but probably occurs in the 
higher subzones of the Upper Gault which have not yet been studied. The closest 
species of M. longispina seems to be M. parallela Alexander 1934 from the Santonian 
of Texas which is a little smaller and more strongly inflated. Good details of 
M. parallela are lacking and it is possible that the two forms are synonymous. M. 
longispina differs from M. acanthoptera (Marsson) 1880 in that the latter is much 
smaller and has the spine set much further back on the carapace. The form recorded 
as M. acanthoptera (Marsson) by Alexander (1934) probably belongs to M. parallela. 
As shown earlier M. ^mibonata acanthoptera (Jones & Hinde) is a separate species 
viz. M. umbonatoides Kaye 1964&. 



244 SOME NEW BRITISH ALBIAN OSTRACODA 

Monoceratina sp. 

(PI. II, figs. 9-10) 

Material. A right valve, B.M.N.H. lo. 2945, from the H. spathi Subzone, Middle 
Albian ; Devizes, Wiltshire. 

Measurements. 

Length Height 
(mm.) (m.m.) 

Right valve (B.M.N.H. lo. 2945) . . . . 0-55 0-25 

Description. Valves elongate, with straight parallel dorsal and ventral margins. 
Lateral surface inflated with a vertical median sulcus, limited ventrally by an alate 
expansion. The crest of the ala bears a short broad based conical spine. This spine 
is set just behind the ventral end of the sulcus and is laterally directed. A large 
smooth semicircular swelling occurs dorsally on the antero-lateral surface whilst a 
short slightly curved ridge runs near to the dorsal margin on the postero-lateral 
surface. A weak anterior marginal rib occurs which is not joined to the ala ventrally. 
The ventral and posterior marginal areas are flattened. The lateral surface is smooth. 

Remarks. Only a single specimen of this species has so far been found, which 
seems to bear no relationship to other species of the genus found in the Albian and 
occurs earlier than them (in the H. spathi Subzone). Its closest relative is M. 
bonnetnai Kaye 1964& from which it differs in size and ornamentation and in having 
only one laterally directed spine. 



Family PROGONOCYTHERIDAE 

Genus ACROCYTHERE Neale i960 
Acrocythere striata sp. nov. 
(PI. 4, figs. 4-10) 

Derivation of name. Striatus L. = striate. 

Diagnosis. Acrocythere with ornamentation of numerous longitudinal striate 
ridges. 

Holotype. a left valve, B.M.N.H. lo. 2853, from the basal Upper Gault (Upper 
Albian) ; Pinhay, Devon. 

Paratypes. B.M.N.H. lo. 2851-52, 2854-57. Eight valves and two carapaces 
from the same horizon and locality. 
Measurements. 

Length Height 
(m.m.) (m.m.) 

Left valve (B.M.N.H. lo. 2853, holotype) . . 0-58 0-28 

Right valve (B.M.N.H. lo. 2852) . . . . 0-58 0-23 



SOME NEW BRITISH ALBIAN OSTRACODA 245 

Description. Valves small, elongate, strongly compressed laterally. Dorsal and 
ventral margins long and straight ; tapering slightly posteriorly in the left valves but 
parallel in the right valves. Dorsal margin without marked cardinal angles. Anterior 
margin broadly rounded ; posterior margin pointed at mid-height. Greatest height 
at 5 length ; greatest width at mid-length. Lateral surface ornamented by numerous 
longitudinal striate ridges. The ridges continue over the whole of the lateral surface 
being usually ten in niunber. Anteriorly they tend to swing slightly ventrally and 
certain of them coalesce. Vertical cross ribbing is absent, the area between the 
ridges being finely pitted. Eye spots are absent. 

Duplicature fairly broad crossed by few, very fine, simple radial pore canals (8-10 
anteriorly). Normal pore canals rather few, confined to the crests of the ridges. 
Inner margin and line of concrescence coincide throughout. Muscle scars not seen. 
Hinge strong merodont consisting in the left valves of two deep strongly divided 
terminal sockets separated by a long, straight, coarsely denticulate bar. In the right 
valve there are two triangular terminal teeth, highest away from the centre of the 
valve and each divided into four large denticles. Between the terminal teeth there is 
a long, straight coarsely crenulate (almost interdentate) furrow. 

Remarks. A . striata differs considerably from the other members of the genus in 
ornamentation and lack of even a rudimentary median sulcus and eye spots. Its 
dissimilarities are such that it could possibly belong to a new genus. The hinge and 
basic longitudinal striate ornamentation are strongly reminiscent of Pleurocythere 
and Lophocythere. 

Genus NEOCYTHERE Mertens 1956 
Subgenus PHYSOCYTHERE Kaye 1963 
Neocythere (Physocythere) tenuis sp. nov. 
(PL 6, figs. 14-17) 
Derivation of name. Tenuis L. = thin. 

Diagnosis. Small Neocythere with thin shell and weakly developed hinge. 
Lateral surface devoid of ornamentation. 

HoLOTYPE. A left valve, B.M.N.H. lo. 2889, from Bed i, A. intermedius Subzone, 
Lower Gault ; Small Dole, Henfield, Sussex. 

Paratypes. B.M.N.H. lo. 2890-93 from the same horizon and locality. 

Measurements. 

Length Height 
(mm.) (mm.) 

Left valve (B.M.N.H. lo. 2889 holotype) . . . 0-50 0-30 

Right valve (B.M.N.H. lo. 2890) . . . . 0-50 0-28 

Description. Valves small, ovate, thin shelled. Dorsal margin straight ; 
ventral margin weakly convex and subparallel to it. Anterior and posterior margins 
broadly rounded. Lateral surface smooth, inflated. Greatest height at \ length ; 



246 SOME NEW BRITISH ALBIAN OSTRACODA 

greatest width at mid-length. In dorsal view valves convex with acute anterior and 
posterior ends. A slight ventro-lateral tumidity occurs. Duplicature rather narrow, 
crossed by few (7-8 anteriorly) thick, straight radial pore canals. Inner margin and 
line of concrescence coincide throughout. Normal pore canals rather large, irregularly 
scattered over most of the lateral surface, but forming two concentric rows along the 
crest of the ventro-lateral expansion. Hinge weak having, in the left valves, two 
faintly divided sockets separated by a weakly denticulate bar. Above the median 
bar is a prominent marginal shelf but no accommodation groove. The terminal 
teeth in the right valve are low, triangular in shape and weakly divided. Muscle 
scars a slightly postero-dorsally inclined row of four oval scars with a U-shaped scar 
antero-dorsally and a small oval scar antero-ventrally of them. 

Remarks. This species has been recorded from the Lower Gault A. intermedius 
Subzone at Heniield but is known to occur abundantly in the Upper Gault in East 
Anglia. It differs from other species of Neocythere s.l. in its shape, thinness of shell, 
weak hinge and lack of ornamentation. In shape and size it is nearest to N. (P.) 
pustiilosa Kaye 1965a from the Upper Aptian of the Isle of Wight but lacks the 
strong ornamentation. Certain related undescribed specimens occur in the Aptian 
but they are larger and of unequal inflation. 



Family PROTOCYTHERIDAE 

Genus VEENIA Butler & Jones 1957 
Veenia compressa Kaye 

(PL II, figs. 13-15) 
1965a Veenia compressa Ka.ye : 44, pi. 7, figs. 6, 7. 

Material. Two valves and one carapace, B.M.N.H. lo. 2868-70, from the basal 
Upper Gault ; Pinhay, Devon. 

Remarks. This species previously recorded from the Upper Aptian of the Isle of 
Wight has now been found rather more abundantly and somewhat better preserved 
in the basal Upper Aptian at Pinhay in Devon. These latter specimens are identical 
although somewhat smaller (0-58 mm.). 

Veenia florentinensis Damotte 

(PL II, figs. 1-8) 

1961 Veenia {Protoveenia) florentinensis Damotte : 102, pi. i, figs. 1—3, pi. 2, figs. 1-6. 

Material. B.M.N.H. lo. 2974-75, two valves from the H. spathi Subzone Lower 
Gault ; Henfield. B.M.N.H. lo. 2971-73, 2976-79, ten valves and two carapaces 
from the H. orbignyi Subzone Upper Gault ; Pinhay, Devon. 



S O:\IE NEW BRITISH 



Measurements. 



ALBIAN 


OSTRACODA 




Length 


Height 


(mm,) 


(mm.) 


0-50 


0-28 








0-49 


0-25 








0-70 


0-36 








0-65 


0-35 








0-58 


0-28 








0-54 


0-28 








0-50 


0-30 








0-48 


0-27 








0-48 


0-25 



247 



Left valve (B.M.N.H. lo. 2975) 
Right valve (B.M.N.H. lo. 2974) 
Left valve (B.M.N.H. lo. 2978) 
Left valve (B.M.N.H. lo. 2971) 
Right valve (B.M.N.H. lo. 2976) 
Right valve (B.M.N.H. lo. 2977) 
Left valve (B.M.N.H. lo. 2972) 
Left valve (B.M.N.H. lo. 2979) 
Right valve (B.M.N.H. lo. 2973) 

Remarks. Damotte erected this species in 1961 placing it within a new subgenus 
Protoveenia. This subgenus was differentiated primarily on a basis of having very 
few radial pore canals. Damotte's specimens came from the Lower Gault and 
comparable specimens occur in the H. spathi and A. intermedins Subzones in Britain. 
Other specimens of this species occur at higher horizons particularly in Devon, Dorset 
and the Isle of Wight in clays from the H. orbignyi Subzone, Upper Gault and these 
specimens have a much larger size range than those from the Lower Gault. The 
Lower Gault forms have a length in the range of o-40-o-5o mm., whilst specimens 
from the Upper Gault reach as much as 0-70 mm. in length. The largest specimens 
differ slightly from the smaller ones but specimens of typical Lower Gault size and 
features and all intermediates occur together with them in the same sample. The 
larger specimens are more inflated, have the longitudinal ribs less keel-like than the 
small ones and have the dorsal ridge and antero-dorsal hinge ear less well separated. 
They also have more radial pore canals and therefore make the subgenus Protoveenia 
unusable and its postulated ancestry to Veenia improbable. The smaller specimens 
in the L'pper Gault are identical with Damotte's Lower Gault foims and it appears 
that the species is therefore not only of variable size but that the number of radial 
pore canals is a direct function of the size. Sexual dimorphism is not known from the 
Lower Gault specimens but is well marked in the Upper Gault forms. 



Family TRACHYLEBERIDIDAE 

Genus CYTHEREIS Jones 1849 
Cythereis angulatoides nom. nov. 

1964a. Cytheveis angulata Kaye : 327, pi. 54, fig. 11. 

Remarks. Professor W. A. Van den Bold of Louisiana State University has 
kindly pointed out that the form erected as C. angulata from the Upper Aptian of 
Surrey has the name preocupied by Sars (1866 : 40). I have therefore renamed my 
form C. angulatoides. 



Geol. II 5 



24 



248 SOME NEW BRITISH ALBIAN OSTRACODA 

Cythereis gatyensis Damotte & Grosdidier 

(pl. II, figs. II, 12) 

1963 Cythereis ? gatyensis Damotte & Grosdidier : 58, pi. 3, figs. Sa-g-. 
19636 Cythereis lamplnghi Kaye : 236, pi. 19, figs. 14-16. 
1965a Cythereis lamplitghi Kaj^e, Kaye: 46, pi. 7, figs. 14, 15. 

Material. B.M.N.H. Io. 2966-67. Two specimens from the H. spathi Subzone, 
Lower Gault ; Culham, Oxfordshire. 

Remarks. Specimens kindly sent to me by Dr. E. Grosdidier show that Cythereis 
lamplughi Kaye 19636 is synonymous with Cythereis gatyensis Damotte & Grosdidier 
1963, the latter having two months' priorit}'. C. gatyensis is abundant but restricted 
to the H. spathi Subzone in the Gault. It is also known from the Upper Aptian of 
the Isle of Wight. It is a valuable index fossil for the H. spathi Subzone having been 
found in clays of that age at Speeton, West Heslerton, Culham and Henfield. 

Cythereis glabrella Triebel 

(PI. 10, figs. 5-8) 
1940 Cythereis glabrella Triebel : 196, pi. 6, figs. 60-62. 

Material. B.M.N.H. Io. 2962-63. Two valves from the basal Upper Gault at 
Pinhay, Devon. 
Measurements. 

Length Height 

(mm.) (mm.) 

Left valve (B.M.N.H. Io. 2962) . . . . 0-90 0-55 

Right valve (B.M.N.H. Io. 2963) .... 0-90 0-50 

Remarks. This smooth, inflated species is not common in the British Gault 
having been found so far only at Pinhay and in the top Red Chalk at Speeton, Yorks. 
It differs from the much more abundant C.folkstonensis Kaye 1964& in its less angular 
appearance, smaller size and in having a smooth rather than spined median rib. It is 
similar to C. niida Jones & Hinde i8go differing in being larger, more inflated and in 
having the median rib well developed. 

Cythereis pinhayensis sp. nov. 

(PI. 9, figs. 1-8) 

Derivation of name. After Pinhay Point, Devon, the only known occurrence of 
the species. 

Diagnosis. Small, reticulate but not spined Cythereis with prominent muscle node 
and weakl}^ convergent long margins. 

HoLOTYPE. A male left valve, B.M.N.H. Io. 2946, from the basal Upper Gault ; 
Pinhav Point, Devon. 



SOME NEW BRITISH ALBIAN OSTRACODA 249 

Paratypes. B.M.N.H. To. 2947-54. Ten valves and two carapaces from the 
same horizon and locaht}'. 
Measurements. 

Length Height 

(mm.) (mm.) 

Maleleft valve (B.M.N.H. lo. 2946 holotype) . . 0-95 0-50 

Male right valve (B.M.N.H. lo. 2947) . . . 0-95 0-48 

Female left valve (B.M.N.H. To. 2948) . . . 0-85 0-50 

Description. Valves elongate, laterally compressed. Dorsal and ventral margins 
straight, converging slightly posteriorly. Antero-dorsal and postero-dorsal cardinal 
angles well marked. Anterior margin broadly rounded, posterior triangular, angled 
just below mid-height. Anterior and posterior margins strongly denticulate, each 
tubercle marking the distal end of a radial pore canal. Greatest height at J length ; 
greatest width at f length. Eye tubercle glassy, distinct, joined to a prominent 
anterior marginal ridge. This ridge bears a row of tubercles on its upper surface 
each marking the extremities of a pseudoradial pore canal. A large, high, smooth 
subcentral muscle node occurs, separated from the anterior end of a short median 
longitudinal ridge but it is joined by a weak vertical ridge to the eye tubercle. A 
short straight low ridge follows the dorsal margin, being connected by a vertical cross 
ridge posteriorly to the posterior end of the median ridge. A shallow depression 
separates the anterior end of the dorsal rib and the muscle node. A prominent ridge, 
increasing in height posteriorly follows the ventral margin. Anteriorly it is con- 
tinuous with the anterior marginal ridge. A small tubercle, the porenkagel, occurs at 
mid-length between the median and ventral longitudinal ribs. Ventral undersurface 
covered with faint longitudinal riblets. Lateral surface strongly reticulate. 

Duplicature fairly broad, crossed by numerous thin, straight radial and pseudo- 
radial pore canals. Hinge strong amphidont, consisting in the left valve of two 
terminal divided sockets, open ventrally, separated by a long, straight, smooth bar. 
Below the anterior end of the median bar is a high, smooth, circular tooth whilst 
above the median element is a narrow marginal shelf. In the right valve there are 
two high boss-like terminal teeth, each with weak lobation on the summit, separated 
by a long, straight, shelf-like median furrow. Anteriorly the furrow opens into a deep 
circular socket. Above the median furrow is a high narrow smooth marginal bar. 

Remarks. C. pinhayensis is only known from the basal Upper Gault at Pinhay 
Point in Devon where it is one of the most abundant members of the fauna. It 
differs from the C. reticulata (Jones & Hinde) 1890, C. hirsuta Damotte & Grosdidier 
1963 group in being smaller, less inflated and lacking the spination of the longitudinal 
ribs. C. lurmannae Triebel 1940a and C. corrigenda Kaye 19646 differ in being more 
strongly compressed laterally, more strongly convergent posteriorly and lacking the 
pronounced reticulation. C. glabrella Triebel 1940a is more inflated and smooth 
whilst C. folkstonensis Kaye 19646 has all the characteristics of C. reticulata in 
addition to being smooth. C. matronae Damotte & Grosdidier 1963 is smaller, smooth 
and has rows of spines rather than well marked longitudinal ridges whilst Cythereis 
thorenensis 1940* Triebel lacks the muscle node and median longitudinal ridge. The 



250 SOME NEW BRITISH ALBIAN OSTRACODA 

closest form is C. geometrica fittoni Kaye ig65a from the Upper Aptian of the Isle of 
Wight which differs in being more angular in outline, particularly posteriorly and in 
having the long margins more strongly convergent. True C . geometrica Damotte & 
Grosdidier 1963^ is smooth whilst C. sutterhvensis Kaye 1965ft and C. heknmensis 
Triebel 1940(2 have less marked reticulation and a short spined median longitudinal 
rib. C. acuticostata Triebel 1940a has the median rib and muscle node joined. C. 
hlanda Kaye 19636 is similar and lacks surface reticulation. C. angtdata Kaye 1964a 
differs in the distribution of the ribs and lacks a muscle node whilst C. cristata Kaye 
1964^ is much more strongly compressed, pitted and has weak ribbing. 

Suborder MYODOCOPINA 

Family CONCHOECIIDAE 

Genus CONCHOECIA Dana 1849 

Conchoecia sp, a 

(PI. 2. figs. 2,7) 

Material. A single carapace, B.M.N.H. lo. 2833, from the Upper Albian, 
H. orbignyi Subzone ; Wrotham, Kent. 

Measurements. 

Length Height 
(mm.) (mm.) 
Carapace (B.M.N.H. lo. 2833) 0-58 0-35 

Remarks. Only one previous fossil occurrence of the genus Conchoecia is known. 
This species, C. cretacea Pokorny 1964, is known as two pyrite infilled carapaces from 
the Upper Cretaceous of Bohemia. The present record extends the range of the 
genus into the Lower Cretaceous. The valves are extremelv thin and very fragile, 
the single specimen being a pyrite filled carapace. It is similar to Pokorny's form in 
shape and ornamentation but is approximately half the size. This may indicate that 
it is an instar of C. cretacea. The anterior rostrum is particularly well developed but 
the posterior prolongation of Pokorny's form is absent. It also differs in ornamenta- 
tion being less strongly ribbed and having the cross ribs more prominent. 

Conchoecia sp. A. differs markedly from the form described as ? Conchoecia sp. B. 
in shape, ornament and in the weak or absent rostrum in the latter. 

? Conchoecia sp. b 
(PI. I, figs. 1-2) 

Material. A carapace, B.M.N.H. lo. 2819, from the basal Upper Albian, 
H. orbignyi Subzone ; Wrotham, Kent. 

Measurements. 

Length Height 
(mm.) (mm.) 

Carapace (B.M.N.H. lo. 2819) . . . . 0-87 0-62 



SOME NEW BRITISH ALBIAN OSTRACODA 251 

Description. One large pyrite filled carapace of this species has been found 
together with four smaller similarly preserved specimens. The shell is extremel}' 
thin and is broken in places. In shape the specimens are suboval and inflated ; the 
greatest height and width being at f length. The dorsal margin is strongly arched ; 
the ventral margin is weakly convex or straight. The anterior and posterior ends 
are bluntly rounded, without prominent extensions or a marked rostrum. The valves 
are ornamented with a series of shallow longitudinal grooves. They are scattered 
irregularly over the surface, being more concentrated posteriorly. They measure 
0-I0-0-I2 mm. in length. 

Remarks. The inflation, shape and ornamentation of the specimen differs some- 
what from Pokorny's form and it is undoubtedly a distinct species. The lack of a 
pronounced rostrum could place it in a different genus, its similarities in shape and 
ornamentation layout to the Palaeozoic Entomozoidae perhaps indicating a relation- 
ship. 

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252 SOME NEW BRITISH ALBIAN OSTRACODA 

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Milbourne, R. a. 1955. The Gault at Greatness Lake, Sevenoaks. Kent. Proc. Geol. Ass., 

London, 66 : 235-242. 

1961. Field Meeting in the Gault at Small Dole, near Henlield, Sussex, Proc. Geol. Ass., 

London, 72 : 135-138. 

1963. The Gault at Ford Place, Wrotham, Kent, Proc. Geol. Ass., London, 74 : 55-80. 

Moore, R. C. (Editor) ig6i. Treatise on Invertebrate Palaeontology, Q. Ostracoda. xxii, -|- 442 
pp., 334 figs. Kansas. 

Neale, J. W. i960. Marine Lower Cretaceous Ostracoda from Yorks., England. Micro- 
paleontology, New York. 6 : 203-224, pis. 1-4. 

1961. The Senonian (Upper Cretaceous) Ostracod Paracypris siliqua Jones & Hinde 1890, 

Ann. Mag. Nat. Hist., London (13) 4 : 193-197, pi. 7. 

1962. Ostracoda from the type Speeton Clay (Lower Cretaceous) of Yorkshire. Micro- 
paleontology, New York, 8 : 425-484, pis. 1-13. 

Oertli, H. J. 1958. Les Ostracodes de L'Aptien-Albien d'Apt. Rev. Inst, franc. Petrole, 
Paris, 13 : 1499-1537, pis. 1-9. 



SOME NEW BRITISH A LB I AN OSTKACODA 253 

Oertli, H. J. 1959- Eiiryitycythere und Parexophthalmocythere zwei neue Ostrakoden- — 
Gattungen aus der Unterkreide Westeuropas. Paldovt. Z., Stuttgart, 33 : 241-246, pi. 32. 

Owen, H. G. 1958. Lower Gault Sections in the Northern Weald and the Zoning of the Lower 
Gault. Proc. Geo!. Ass., London, 69 : 148-165. 

1963. Some Sections in the Lower Gault of the Weald. Proc. Geol. Ass., London, 74 .' 

35-54- 
PoKORNY, V. 1964. Conchoecia ? cvetacea n. sp., first fossil species of the family Halocy- 

prididae (Ostracoda Crustacea) Acta. Univ. Carolinae Geol., Prague, 2 : 175-180, pi. i. 
Reuss, a. E. 1845-6. Die Versteinerungen der Bohmische Kreideformation, 1 : 1-58, pis. 1-13. 

2 : 148, pis. 14-51. 
1874. Die Foraminiferen, Bryozoen und Ostracoden des Planers 3. Die Ostracoden des 

Saachsuchen Planers. Palaeontographica, Stuttgart, 20 : 138-154, pis. 26-28. 
SzczECHURA, J. 1964. Monoceratina Roth (Ostracoda) from the Upper Cretaceous and Lower 

Palaeocene of North and Central Poland. Acta. Pal. polonica, Warsaw, 9 : 357-406, pis. 

i-ii. 
Triebel, E. 1938. Ostracoden Untersuchungen. i. Protocythere und Exophthalmocythere, 

Zwei Neue Ostracoden — Gattungen aus der Duetschen Kreide. Senckenbergiana, Frank- 
furt. a.M., 20 : 178-200, pis. 1-3. 
1938a. Die Ostracoden der Deutschen Kreide XL Die Cytheridea Arten der Untern Kreide. 

Senckenbergiana, Frankfurt a.M., 20 : 471-501, pis. 1-6. 
1940. Die Ostracoden der Deutschen Kreide III Cytherideinae und Cytherinae aus der 

Unteren Kreide. Senckenbergiana, Frankfurt a.M., 22 : 160-227, pis. i-io. 
1941. Zur Morphology und Okologie der Fossilen Ostracoden, mit Beschreibung einigen 

Neuer Gattungen und Arten. Senckenbergiana, Frankfurt a.M., 23 : 294-400, pis. 1-15. 
Veen, J. E. 1936. Die Cytheridae der Mastrichter Tuffkreide und des Kunrader Korallen 

Kalkes von Sud-Limburg III Die Gattungen Loxoconcha, Monoceratina, Paracytheridea, 

Xesteloberis, Cytheropteron und Cytherura. Natiiurh. Maandbl., Maastricht, 25 : 21-113, 

pis. 1-4. 



PLATE I 

All figures x 60 

? Conchoecia sp. B p. 250 

Fig. I. Carapace, from left. lo. 2819, Wrotham 
Fig. 2. Carapace, from right. lo. 2819, Wrotham 



Fig. 
Fig. 
Fig. 
Fig. 
Fig. 



Fig. 8. 
Fig. 9. 
Fig. 10. 
Fig. II. 
Fig. 12. 



Monoceratina longispina (Bosquet) p. 242 

Adult left valve, lateral view. lo. 2820, Wrotham 
Adult right valve, lateral view lo. 2821, Wrotham 
Juvenile carapace, lateral view lo. 2822, Wrotham 
Juvenile carapace, dorsal view. lo. 2822, Wrotham 
Juvenile left valve, lateral view. I0.2823, Wrotham 

Clithrocytheridea heslertonensis Kaye p. 228 

Female left valve, lateral view. lo. 2825, W. Heslerton 
Female right valve, lateral view. lo. 2826, W. Heslerton 
Female right valve, internal view. lo. 2826. W. Heslerton 
Female carapace, dorsal view. lo. 2827, W. Heslerton 
Female left valve, internal view. lo. 2825, W. Heslerton 



BuIl.B.M. [K.H.) Geo!. 11,5 



PLATE 1 




PLATE 2 

All figures x 60 

Bairdia pseudoseptentrionalis (IVIertens) p. 223 

Fig. I. Left valve, lateral view. lo. 2828, Arlesey 

Fig. 3. Right valve, lateral view. lo. 2929, Arlesey 

Fig. 4. Right valve, internal view. lo. 2830, Arlesey 

Fig. 5. Left valve, internal view. To. 2831, Arlesey 

Fig. 6. Carapace, from right. lo. 2832, Arlesey 

Conchoecia sp. A p. 250 

Fig. 2. Carapace, from left. lo. 2833, Wrotham 
Fig. 7. Carapace, from right. lo. 2833 Wrotham 



Bull.B.M. (N.H.) Geol. 11, 5 



PLATE 2 




Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 7 
Fig. 8 



Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 



9- 

lO. 

II. 

12. 

13- 
14- 



PLATE 3 
All figures x 8o 
Pontocyprella semiquadrata sp. nov. 



p. 224 



Left valve holotype lateral view lo. 2834. Wrotham 
Right valve, dorsal view. lo. 2836, Wrotham 
lo. 2835, Wrotham 
lo. 2836, Wrotham 
lo. 2837, Wrotham 
lo. 2835, Wrotham 
Left valve, holotype, dorsal view. lo. 2834, Wrotham 
Right valve, internal view. lo. 2837, Wrotham 



Left valve, lateral view. 
Right valve, lateral view. 
Right valve, lateral view. 
Left valve, internal view. 



Dolocytheridea typica sp. nov. p. 230 

Right valve, lateral view. lo. 2838, Pinhay 

Left valve, holotype, lateral view. lo. 2839, Pinhay 

Left valve, lateral view. lo. 2840, Pinhay 

Carapace, dorsal view. lo. 2841, Pinhay 

Right valve, lateral view. lo. 2842, Pinhay 

Left valve, internal view. lo. 2843, Pinhay 

Krausella sp. p 227 

Fig. 15. Right valve, lateral view. lo. 2845, Ely 
Fig. 16. Right valve, internal view. lo. 2845, Ely 



Bull.B.M. (N.H.) Geol. 11, 5 



PLATE 3 




PLATE 4 

All figures x 80 

Polycope nuda sp. nov. p. 221 

Fig. I. Left valve, holotype, lateral view. lo. 2847, Wrotham 
Fig. 2. Carapace, from left. lo. 2848, Wrotham 
Fig. 3. Carapace, from left. lo. 2849, Wrotha 



Fig. 4 
Fig. 5 
Fig. 6 
Fig. 7 
Fig. 8 
Fig. 9 



Acrocythere striata sp. nov. p. 244 

Left valve, lateral view. lo. 2851, Pinhay 

Right valve, lateral view. lo. 2852, Pinhay 

Left valve, holotj'pe, lateral view. lo. 2853, Pinhay 

Carapace, dorsal view. lo. 2854, Pinhay 

Right valve, lateral view. lo. 2855, Pinhay 

Right valve, lateral view. lo. 2856, Pinhay 



Fig. id. Right valve, internal view. lo. 2856, Pinhay 

Polycope oweni sp. nov. p. 222 

Fig. II. Carapace, from left. lo. 2858, Wrotham. 

Fig. 12. Left valve, holotype, lateral view. lo. 2859, Wrotham. 

Fig. 13. Left valve, lateral view. lo. 2860, Wrotham. 

Fig. 14. Carapace, from left. lo. 2861, Wrotham 

Fig. 15. Left valve, internal view. lo. 2862, Wrotham 



Bull. B.M. (N.H.) Geol. 11, 5 



PLATE 4 







f* •'i** 






\^ 






14 




PLATE 5 

All figures x 80 

Schuleridea dimorphica sp. nov. p. 228 

Fig. I. Male left valve, holotype, lateral view. lo. 286<j, Wrotham 

Fig. 2. Male left valve, internal view. lo. 2865, Wrotham 

Fig. 3. Male right valve, lateral view. lo. 2866, Wrotham 

Fig. 4. Female left valve, lateral view. lo. 2867, Wrotham 

Fig. 5. Female right valve, lateral view. lo. 2868, Wrotham 

Fig. 6. Male right valve, internal view. lo. 2869, Wrotham 

Orthonotacythere fordensis sp. nov. p. 237 

Fig. 7. Male left valve, holotype, lateral view. lo. 2871, Wrotham 

Fig. 8. Male right valve, lateral view. lo. 2872, Wrotham 

Fig. 9. Female left valve, lateral view. lo. 2873, Wrotham 

Fig. id. Male carapace, dorsal view. lo. 2874, Wrotham 

Fig. II. Female right valve, lateral view. lo. 2875, Wrotham 

Fig. 12. Male left valve, internal view. lo. 2876, Wrotham 

Fig. 13. Female right valve, internal view. lo. 2877, Wrotham 



Bull. B.M. (N.H.) Geol. 11,5 



PLATE 5 




12 



13 



PLATE 6 

All figures x 80 

Cy t heropteron (Eocytheropteron), protonsa sp. nov. p. 235 

Fig. I. Left valve, holotype, lateral view. lo. 2879, Wrotham 

Fig. 2. Right valve, lateral-view. lo. 2880, Wrotham 

Fig. 3. Left valve, dorsal view. lo. 2881, Wrotham 

Fig. 4. Left valve, lateral view. lo. 2881, Wrotham 

Fig. 5. Right valve, dorsal view. lo. 2880, Wrotham 

Fig. 6. Left valve, holotype, internal view. lo. 2879, Wrotham 

Habrocythere fragilis Triebel p. 229 

Fig. 7. Male left valve, normal type, lateral view. lo. 2882, Wrotham 

Fig. 8. Male left valve anomalous type, lateral view. lo. 2883, Wrotham 

Fig. 9. Female left valve, normal type, lateral view. lo. 2884, Wrotham 

Fig. 10. Male left valve, anomalous type, lateral view. lo. 2885, Wrotham 

Fig. II. Female right valve, anomalous type, lateral view. lo. 2886, Wrotham 

Fig. 12. Male right valve, normal type, lateral view. lo. 2887, Wrotham 

Fig. 13. Female right valve, normal type, lateral view. lo. 2888, Wrotham 

Neocythere (Physocythere) tenuis sp. nov. p. 245 

Fig. 14. Left valve, holotype, lateral view. lo. 2889, Henfield 

Fig. 15. Right valve, lateral view. lo. 2890, Henfield 

Fig. 16. Left valve, lateral view, lo. 2891, Henfield 

, Fig. 17. Right valve, lateral view. lo. 2892, Henfield 



Bull. B.M. (N.H.) Geol. 11, 5 



PLATE 6 




PLATE 7 

All figures x 80 

Orthonotacythere spinifera sp. nov. p. 239 

Fig. I. Right valve, lateral view. lo. 2894, Wrotham 

Fig. 2. Right valve, lateral view. lo. 2896, Henfield 

Fig. 3. Left valve, holotype, lateral view. lo. 2895, Wrotham 

Fig. 5. Left valve, lateral view. lo. 2897, Henfield 



Fig. 4 
Fig. 6 
Fig. 7 
Fig. 8 
Fig. 9 



Cytheropteron (C.) milbournei sp. nov. 



Left valve, holotype, lateral view. lo. 2 
Left valve, lateral view. lo. 2899, Wrotham 
Right valve, lateral view. lo. 2900, Sevenoaks. 
Carapace, dorsal view. lo. 2901, Sevenoaks. 
Left valve, internal view. lo. 2902, Sevenoaks 



P- 233 

i, Wrotham 



Cytheropteron ( Intracytheropteron) obscura sp. nov. p. 236 

Fig. 10. Left valve, holotype, lateral view. lo. 2903, Wrotham 
Fig. II. Right valve, lateral view, I0.2904, Wrotham 
Fig. 12. Right valve, lateral view. lo. 2905, Wrotham 

Cytheropteron (C.) nanissimum nanissimum Damotte & Grosdidier p. 234 



Fig. 13. 
Fig. 15. 



Left valve, lateral view. 
Left valve, lateral view. 



lo. 2907, Wrotham 
lo. 2908, Wrotham. 



Cytheropteron (C.) nanissimum Jenestrata ssp. nov. p. 234 

Fig. 14. Left valve, lateral view. lo. 2909, Henfield 

Fig. 16. Left valve, holotype, lateral view. lo. 2910, Henfield 

Fig. 19. Left valve, lateral view. lo. 2911, Henfield 



Eucytherura afi. nuda Kaye p. 231 



Fig. 17. Left valve, lateral view. 
Fig. 18. Right valve, dorsal view. 



lo. 2912, Wrotham 
lo. 2913, Wrotham 



Fig. 


20. 


Fig. 


21. 


Fig. 


22. 


Fig. 


23- 


Fig. 


24 


Fig. 


25- 



Argilloecia valvula sp. nov. 



p. 225 



Right valve, holotype, lateral view. lo. 2914, Wrotham 
Left valve, lateral view. lo. 2915, Wrotham 
Carapace, from left. lo. 2916, Wrotham 
Right valve, lateral view., lo. 2917, Wrotham 
Left valve, lateral view. lo. 2918, Wrotham 
Right valve, internal view. lo. 2919, Wrotham 



Bull.B.M. L\.H.) Geol. 11, 5 



PLATE 7 




^Ib 





PLATE 8 

All figures x 120 

Hemicytherura euglyphea sp. nov. p. 231 

Fig. I. Male left valve, holotype, lateral view. lo. 2921, Wrotham 

Fig. 2. Female right valve, lateral view. lo. 2922, Wrotham 

Fig. 3. Male right valve, lateral view. lo. 2923, Wrotham 

Fig. 4. Female left valve, lateral view. lo. 2924, Wrotham 

Or thonotacy there tninutissima sp. nov. p. 239 

Left valve, holotype, lateral view. lo. 2926, Wrotham 
Right valve, lateral view. lo. 2927, Wrotham 
Carapace, dorsal view. lo. 2928, Wrotham 
Left valve, internal view. lo. 2929, Wrotham 
Right valve, lateral view. lo. 2930, Wrotham 
Left valve, lateral view. lo. 2931, Wrotham 
Left valve, lateral view. lo. 2932, Wrotham 

Cytheropteron (C.) arguta sp. nov. p. 232 

Right valve, lateral view. lo. 2933, Wrotham 

Carapace, dorsal view. lo. 2934, Wrotham 

Right valve, lateral view. lo. 2935, Wrotham 

Left valve, holotype, lateral view. lo. 2936, Wrotham 

Left valve, lateral view. lo. 2937, Wrotham 

Left valve, internal view. lo. 2938, Wrotham 



Fig. 


5- 


Fig. 


6. 


Fig. 


7- 


Fig. 


8. 


Fig. 


9- 


Fig. 


10 


Fig. 


II 



Fig. 


12. 


Fig. 


13- 


Fig. 


14- 


Fig. 


15- 


Fig. 


16. 


Fig. 


17- 



Bull. B.M. (X.H.) Geol. 11, 5 



PLATE 8 





^^-1 







Xy 



/I 







IS 






^•^f^^^s 



17 



PLATE 9 



Fig. I 
Fig. 2 
Fig. 3 
Fig. 4 
Fig. 5 
Fig. 6 
Fig. 7 
Fig. 8 



All figures x 6o 

Cythereis pinhayensis sp. nov. p. 248 

Male left valve, holotype, lateral view. lo. 2946, Pinhay 
Male right valve, lateral view. lo. 2947, Pinhay 
Female left valve, lateral view. lo. 2948, Pinhay 
Female right valve, lateral view. lo. 2949, Pinhay 
Male left valve, lateral view. lo. 2950, Pinhay 
Female carapace, dorsal view. lo. 2951, Pinhay 
Female right valve, internal view. lo. 2952, Pinhay 
Male left valve, internal view. lo. 2953, Pinhaj^ 



Paracypris wrothamensis sp. nov. 



226 



Fig. 9. Left valve, internal \iew. lo. 2955, Wrotham 

Fig. 10. I^eft valve, lateral view. lo. 2956, Wrotham 

Fig. II. Carapace, dorsal view. lo. 2957, Wrotham 

Fig. 12. Right valve, lateral view. lo. 2958, Wrotham 

Fig. 13. Left valve, holotype, lateral view. lo. 2959, Wrotham 

Fig. 14. Right valve, lateral view. lo. 2960, Wrotham 



Bull. B.M. {N.H.) Geol. 11, 5 



PLATE 9 




Fig. 


I. 


Fig. 


2. 


Fig. 


3- 


Fig. 


4- 


Fig. 


5- 


Fig. 


6. 


Fig. 


7- 


Fig. 


8. 


Fig. 


Al 

9- 


Fig. 


lO. 



PLATE lo 

All figures X 6o 

Alatacythere robusta langi ssp. nov. p. 241 

Male left valve, holotype, lateral view. lo. 2940, Pinhay 
Female left valve, lateral view. lo. 2941, Pinhay 
Female right valve, lateral view. lo. 2942, Pinhay 
Male right valve, lateral view. lo. 2943, Pinhay 

Cy t hereis glabrella Triehel p. 248 

Left valve, lateral view. lo. 2962, Pinhay 
Left valve, dorsal view. lo. 2962, Pinhay 
Right valve, dorsal view. lo. 2963, Pinhay 
Right valve, lateral view. lo. 2963, Pinhay 

Alatacythere robusta robusta (Jones & Hinde) p. 240 

Right valve, lateral view. lo. 2964, Arlesey 
Left valve, lateral view. lo. 2965, Arlesey 



Bull. B.M. {N.H.) Geol. 11,5 



PLATE 10 





S.%r. 




*-< 



\ 



A 




-^w"^-: 



# 






a 



■y 




10 



PLATE II 

All figures x 80 

Veenia florentinensis Damotte p. 246 

Fig, I. Left valve, lateral view. lo. 2971, Pinhay 

Fig. 2. Left valve lateral view lo. 2971, Pinhay 

Fig. 3. R-ght valve lateral view, lo: 2973 Pinhay 

Fig. 4. Right valve, lateral view. lo. 2974, Henfield 

Fig. 5. Left valve, lateral view. lo. 2975, Henfield 

Fig. 6. Right valve, lateral view. lo. 2976, Pinhay 

Fig. 7. Right valve, lateral view. lo. 2977, Pinhay 

Fig. 8. Left valve, lateral view. lo. 2978, Pinha^' 

Monoceratina sp. p. 244 

Fig. 9. Right valve dorsal view. lo. 2945, Devizes 
Fig. 10. Right valve, lateral view. lo. 2945, Devizes 

Cythereis gatyensis Damotte & Grosdidier p. 248 

Fig. II. Right valve, lateral view. lo. 2966, Culham 
Fig. 12. Left valve, lateral view. lo. 2967, Culham 

Veenia compressa Kaye p. 246 

Fig. 13. Right valve, lateral view. lo. 2968, Pinhay 
Fig. 14. Carapace, dorsal view. lo. 2969, Pinhay 
Fig. 15. Left valve, lateral view. lo. 2970, Pinhay 



Bull. B.M. (N.H.) Geol. 11, 5 



PLATE 11 




i . I 24JAr 

TERTIARY RED ALGAE FROM ^^^^ 
BORNEO 



J. HARLAN JOHNSON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. ii No. 6 

LONDON: 1966 



. i X JAM 19( 

TERTIARY RED ALGAE FROM BORNEO Kit 



BY 



J. HARLAN JOHNSON 

(Professor, Department of Geology, Colorado School of Mines) 



Pp. 255-280 ; 6 Plates ; i Text-figure 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

GEOLOGY Vol. 11 No. 6 

LONDON: 1966 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 6 of the Geological 
[Palaeontological] series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 



Trustees of the British Museum (Natural History) 1966 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 7 January, 1966 Price £1 10s. 



TERTIARY RED ALGAE FROM BORNEO 

By J. HARLAN JOHNSON 





CONTENTS 














Page 


I. 


Acknowledgments ......... 257 


II. 


Introduction 












257 


III. 


Locality and age data . 












258 


IV. 


Systematic descriptions 

Family CORALLINACEAE 
Genus Archaeolithothamnium 
Lithothamnium 
Mesophyllum 
Lithophylliim 
Lithoporella . 
Melobesia 
Dermatolithon 
Thaumatoporella . 
Corallina 
Jania . 
Amphiroa 
Subterraniphyllum 
Distichoplax . 
Family SOLENOPORACEAE 

Genus Solenomeris . 
Family SIPHONOCLADACEAE 
Genus Pycnoporidium 












261 
261 
261 
266 
267 
269 
271 
273 
273 
273 
274 

275 
276 
277 
277 
. 278 
. 278 
278 
278 


V. 


References .... 












279 



SYNOPSIS 
Forty-one species belonging to fifteen genera are described. Of these two species and one 
variety are considered to be new. The material comes from the Melinau Gorge and from the 
Upper Baram and Belukan Rivers, Northeast Sarawak, Borneo, Malaysia. 

I. ACKNOWLEDGMENTS 

I AM deeply indebted to Dr. C. G. Adams for his kindness in arranging for me to 
borrow this collection from the British Museum (Natural History) for study and for 
his assistance in supplying necessary data. Thanks are due the Colorado School of 
Mines who supplied the working space and laboratory facilities, and the Colorado 
School of Mines Foundation, Inc., for financing the project. Our secretary Mrs. 
Ruth Loomis typed the manuscript and read the proof. 

II. INTRODUCTION 

The coralline algae of the Paleocene are only slightly known. As of October 1963 
only a single short paper had been published on them. In 1961 the author started a 
series of studies of the red calcareous algae from the Paleocene based on a series of 
collections obtained from oil companies, geological surveys, and museums. This 
paper, in so far as it deals with Paleocene algae from the Upper Baram area of 
Sarawak, represents the third of these studies. It does, however, also deal with 



GEOL. II, 6 



25 



258 



TERTIARY RED ALGAE FROM BORNEO 



Miocene algae from the Upper Baram and with Upper Eocene (Tb) , Ohgocene (Ted) , 
and Lower Miocene (Lower and Upper Te) algae from the Melinau limestone in north- 
east Sarawak. This study is based on a collection of petrographic slides belonging 
to the British Museum (Natural History). They were made from a series of lime- 
stone samples collected by the Geological Survey of Sarawak, Sarawak Shell Oilfields 
Ltd., and by Dr. C. G. Adams who has also studied the foraminifera of these limestones 
and has supplied the data for determining the age of the samples. 



III. LOCALITY AND AGE DATA 

Limestones from the Upper Baram and Belukan Rivers 

The locations of the samples from this area are shown on Text-fig. i. The precise 
stratigraphical relationships of most samples are uncertain since the geological 
structure of the district is only known in outhne. This is particularly true of the 
outcrops along the Belukan River where the tectonic picture appears to be fairly 
complex. The geological setting of the major limestones in the area has been briefly 
described by Liechti et al. (i960). All the Paleocene limestones in the area were 
assigned to the Kelalan Formation by Haile (1962). The same author considered 
that the associated Miocene Umestones should be assigned to the Melinau Limestone 
Formation. 

It should be noted that some of the samples dated as Paleocene are from conglom- 
eratic limestones, and the possibility that the fdssil content of these samples is 
reworked cannot be entirely ignored. All such samples are asterisked in the list 
below. 



Limestones from the Upper Baram and Belukan Rivers 



Sample 
S-7058 

S-7059 
S-7063 
S-7067 

*S-7o68 
S-7070 
S-7072 

*S-7073 

S-7075 
S-7076 
*S-7077 
S-7080 
S-7081 
S-7082 
S-7085 
S-7087 
S-7095 
S-7111 



number 
(i slide) 
(2 slides) 
(7 slides) 
(i slide) 
(2 slides) 
(i slide) 
(i shde) 
(2 slides) 
(4 slides) 
(i slide) 
(5 slides) 
(3 slides) 
(2 slides) 
(3 slides) 
(i shde) 
(i slide) 
(i slide) 
(2 sUdes) 



Age 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Palaeocene . 
Miocene (Tei-4) 
Miocene (Tei-4) 
Miocene (Tei-4) 
Miocene (Tei-4) 
Palaeocene . 
Palaeocene . 
Miocene (Te) 



Baram River 
Baram River 
Baram River 
Baram River 
Baram River 
Baram River 
Baram River 
Baram River 
Baram River 
Baram River 
Baram River 
Belukan River 
Belukan River 
Belukan River 
Belukan River 
Belukan River 
Kalai River 
Belukan River 



TERTIARY RED ALGAE FROM BORNEO 



259 




26o 



TERTIARY RED ALGAE FROM BORNEO 



Limestones from the Upper Baram and Belukan Rivers fcontd.J 
Sample number Age 

S-7109 (3 slides) . . Miocene (Te) . . Belukan River 

Palaeocene . . . Baram River 

Palaeocene . . . Baram River 

Palaeocene . . . Baram River 

Palaeocene . . . Baram River 

Palaeocene . . . Batu Asi ridge 



*S-6i75 (i slide) 

*S-6i76 (i slide) 

*S-6i77 (i slide) 

BM 47 (i slide) 

BM 13 (i slide) 



The Melinau limestone 

The slides studied are listed below with their age as determined from the fora- 
minifera. Full information regarding the provenance of the samples from the 
Melinau limestone has been given by Adams (1965). It is sufficient to state here that 
this limestone has a maximum thickness of almost 7,000 feet, that its base is in the 
Upper Eocene (Tb) , and that it includes a great thickness of Oligocene (Tc and Td) 
and Lower Miocene (Te 1-5) beds. Most of the Melinau samples referred to in this 
account come from the north face of the Melinau Gorge where a large niunber of 
samples (S-10000-S-10194) were collected in stratigraphical order from the base 
upwards. The remainder come from different parts of the outcrop. Examination 
of the foraminifera has shown that the main stratigraphical boundaries occur at 
about the following levels in the Melinau Gorge : 

Tb/Tc . between S-10085 and S-10094 ; i.e., at least 1,850 feet above the 

base. 
Tc/Td . between S-10152 and S-10153 ; i.e., about 3,290 feet above the base. 
Td/Te . between S-10165 and S-10166 ; i.e., about 3,400 feet above the base. 
The junction between Lower Te (Te 1-4) and Upper Te (Te 5) is not reached 

in the gorge. 

Melinau Limestone 



Sample number 










Age 


S-6610 (2 slides) ..... Oligocene (Tc) 


S-6613 (3 slides) 










Miocene (Te 1-4) 


S-6614 (3 slides) 










Miocene (Te 5) 


S-6624 (2 slides) 










Miocene (Te 5) 


S-6921 (3 slides) 










Oligocene (Tc) 


S-6952 (3 slides) 










Oligocene (Tc) 


S-6970 (i sHde) 










Miocene (Te 5) 


S-6971 (i sHde) 










Miocene (Te 5) 


S-6972 (i shde) 










Miocene (Te 5) 


S-6976 (i shde) 










Miocene (Te 5) 


S-6977 (2 slides) 










Miocene (Te 5) 


S-6979 (i slide) 










Miocene (Te 5) 


S-6620 (i slide) 










Oligocene (Te 1-4) 


S-6623 (2 slides) 










Oligocene (Te 1-4) 



TERTIARY RED ALGAE FROM BORNEO 



261 



BRITISH MUSEUM SLIDES 
GEOLOGICAL SURVE Y— S AR AWA K 



Samples from the Melinau Gorge 



Sample number 


Age 


S-iooo6a 


I slide) 


Eocene (Tb) 


S-iooi5b 


[i slide) 


Eocene (Tb) 


S-ioo2ia ( 


'i slide) 


Eocene (Tb) 


S-ioo25a ( 


I slide) 


Eocene (Tb) 


S-ioo3oa 


I slide) 


Eocene (Tb) 


S-ioo32a 


I slide) 


Eocene (Tb) 


S-ioo4oa 


'i slide) 


Eocene (Tb) 


S-ioo55a ( 


2 slides) 


Eocene (Tb) 


S-ioo6ob 


I slide) 


Eocene (Tb) 


S-ioo6ia 


2 slides) 


Eocene (Tb) 


S-ioo87b 


I slide) 


? Eocene (?Tb) 


S-I0096a 


I slide) 


Oligocene (Tc) 


S-ioiooa 


[i slide) 


Oligocene (Tc) 


S-ioioia 


I slide) 


Oligocene (Tc) 


S-ioioga 


[i slide) 


Oligocene (Tc) 



Sample number 
S-ioi45b (i slide) 
S-ioi48b (i slide) 
S-ioi49a (i slide) 
S-10153C (6 slides) 
S-ioi54f (2 slides) 
S-ioi55b (i slide) 
S-ioi6ob (3 slides) 
S-ioi62a (i slide) 
S 10176b (i slide) 
S-ioi77b (i slide) 
S-ioi79a (i slide) 
S-ioi84a (i slide) 
S-ioiSga (i slide) 
S-ioi94b (i slide) 
S-i0207b (i slide) 



1883 
1891 
1961 



IV. SYSTEMATIC DESCRIPTIONS 

Phylum RHODOPHYCOPHYTA Papenfuss 1946 

Class RHODOPHYCEAE Ruprecht 1851 

Order CRYPTONEMIALES Schmitz in Engler 1892 

Family GORALLINACEAE (Lamouroux) Harvey 1849 

Subfamily MELOBESIEAE 

Genus ARCHAEOLITHOTHAMNIUM Rothpletz 1891 

Archaeolithothamnium aschersoni (Schwager) 

(PI. I, figs. 3, 4) 

Lithothamnium aschersoni Schwager : 147, pi. 29, fig. 25. 
Lithothamnium aschersoni (Schwager) Rothpletz : 316. 
Archaeolithothamnium aschersoni (Schwager) Segonzac : 437. 



Age 
Oligocene (Tc) 
Oligocene (Tc) 
Oligocene (Tc) 
Oligocene (Td) 
Oligocene (Td) 
Oligocene (Td) 
Oligocene (Td) 
Oligocene (Td) 
Miocene (Te 1-4) 
Miocene (Te 1-4) 
Miocene (Te 1-4) 
Miocene (Te 1-4) 
Miocene (Te 1-4) 
Miocene (Te 1-4) 
Eocene (Tb) 
South face of gorge 



Description. A crustose form, commonly with small mammillated protuber- 
ances. Hypothallus poorly developed or absent. Hypothallic cells measure 
12-20 [i X 8-12 [I. The perithallus forms most of the crust. Tissue quite regular. 
Cell threads fairly prominent, cross partitions thinner, cells 9-19 n X 8-15 /*. 
Sporangia ovoid, arranged in regular rows (layers), height 73-86 [i, diameter 40-52 fi. 

Remarks. This widespread species is one of the most abundantly represented 
in the Borneo collection. 

Sample number and locality. S-7068, Upper Baram River. 

Age. Paleocene (the figured specimens occur in a conglomeratic limestone). 



262 TERTIARY RED ALGAE FROM BORNEO 

Archaeolithothatnnium cf. cyrenaicum Raineri 

(PI. I, fig. 5) 
1923 Archaeolithothamnium cyrenaicum Raineri : 30, text-fig. 2. 

Description. Thallus crustose with protuberances or short stubby branches. 
PerithaUic tissue fairly regular with well defined cell " layers ". At irregular 
intervals " layers " of very short cells are interlay ered with the normal perithallic 
cells. Normal cells measure 14-30 /i x 8-13 /i. Sporangia in layers or lenses. 
They measure 38-45 /i x 70-74 fi. This is very close to, if not identical with, 
Raineri's species from the Miocene of Cyrenaica. 

Remarks. Represented in the Borneo collection by a single, oblique section. 

Sample number and locality. S-10153, Mehnau Gorge, north-east Sarawak. 

Age. Middle Oligocene (Td). 

Archaeolithothamnium intermedium Raineri 

(PI. I, fig. 6) 

1923 Archaeolithothamnium intermedium Raineri : 29, text-fig. i. 

Description. Crustose with rounded protuberances. The tissue of the pro- 
tuberances is quite regular with " layers " of cells. The cells measure 12-19 /* X 8- 
15 /I. Sporangia moderately to closely packed in regular layers. They measure 
36-42 [I in diameter and 79-1 11 fi high. 

Remarks. This closely approximates Raineri's species. Represented in the 
Borneo collection by a single good specimen and a couple of fragments. 

Sample number and locality. S-7111, Belukan River. 

Age. Lower Miocene (Lower Te). 

Archaeolithothamnium lauensum Johnson & Ferris 
(PL I, fig. I) 
1950 Archeolithothamnium lauensum Johnson & Ferris : 11, pi. i, figs. A, D. 

Description. Thallus forms a fairly regular crust up to 9 mm. thick. Hypo- 
thallus of curved cell threads. Cells square or rectangular in section, measuring 
8-17 II X 8-12 /x. Perithallus compact, quite regular, with cells measuring 
8-15 fi X 8-11 [X. Conceptacles large, commonly much higher than wide, ovoid, 
95-108 n in diameter and 120-205 f^ high. 

Remarks. Represented by only a few specimens, mostly small fragments. 
They fit the description of A. lauensum in all respects. 

Sample number and locality. S-7111, Belukan River. 

Age. Lower Miocene (Lower Te). 



TERTIARY RED ALGAE FROM BORNEO 263 

Archaeolithothamnium liberum Lemoine 

1939 Archaeolithothamnium liherum Lemoine : 61, pL i, fig. 14, text-fig. 26. 

Description. Thallus develops as thin crust from which protuberances may 
arise. Hypothallus consists of curved threads of cells. Cell size 7-28 /t x 7-18 /i. 
Perithallus fairly regular, cell size 8-17 /t x 6-18 /i. 

Remarks. Represented in the Borneo collection by several young infertile crusts. 

Sample number and locality. S-7076, Upper Baram River. 

Age. Paleocene. 

Archaeolithothamnium lugeoni Pfender 

(PL I, fig. 2) 

1926 Archaeolithothamnium lugeoni Pfender : 324, pis. 8, 9. 

1935 Archaeolithothamnium lugeoni Pfender ; Miranda : 280. 

1936 Archaeolithothamnium lugeoni Pfender ; Rama Rao & Pia : 35, pi. 4. 
1939 Archaeolithothamnium lugeoni Pfender ; Lemoine : 52-53. 

Description. Thin crusts growing superimposed to form a thick mass. Hypo- 
thallus very thin or absent, difficult to distinguish and measure. Perithallic tissue 
fairly regular with suggestions of layers of cells, but the horizontal partitions not 
all continuous. Perithallic cells 11-16 /t x 8-13 /jl. Sporangia long ovoid to nearly 
subspherical, dimensions 80-104 /* high and 57-58 /* in diameter. 

Remarks. Represented by only a few fragments in the collection studied. 

Sample number and locality. S-7077, Upper Baram River, Sarawak. 

Age. Paleocene. 



Archaeolithothamnium macrosporangium n. sp. 

(PI- 3. fig- 4) 

Diagnosis. Crustose, hypothallus thin, cells 16-36 /^ x 7-14 /* ; perithallus 
regular, cells 9-29 /i x 7-13 /f. Sporangia abundant and unusually long (135-200/^). 

HoLOTYPE. SHde V-51765. 

Sample number and locality. S-6921, mouth of Tukuruk River, Melinau, 
north-east Sarawak. 

Age. Lower Oligocene (Tc). 

Description. Thallus develops as a crust probably with protuberances or small 
mammelons on the upper surface. Several may grow superimposed. Hypothallus 
slightly to moderately developed consisting of curved cell threads. Cells measure 
16-36 [I X 7-14 {i. Perithallus regular with the appearance of cell " layers " as 
well as cell threads. Both the horizontal and vertical partitions between the cells 



264 TERTIARY RED ALGAE FROM BORNEO 

are moderately thick. Cells measure 9-29 fi x 7-13 /^. Sporangia numerous and 
unusually large. Commonly they occur in layers or lenses, appearing as rows in 
vertical sections. Sporangia measure 1 17-124 /i in diameter and 144-200 /i high. 
Where closely packed they are long ovoid in shape. If loosely packed, they tend 
to be round ovoid to nearly spherical. 

Remarks. This species is characterized by the cell size, regular perithallic tissue, 
and unusually large, especially unusually long, sporangia. It belongs in the same 
general group as A. nummuliticum (Giimbel) Rothpletz, A. saipanense Johnson, 
and A. lauensum Johnson & Ferris. It differs from A. nummuliticum in having 
larger, especially longer, cells, better developed hypothallus, more regular perithallic 
tissue, and much larger sporangia. A. saipanense has smaller cells which are nearly 
cubic in the perithallic tissue, and smaller sporangia (50-100 /t x 70-140 [i against 
117-124 // X 144-200 /.i). A. lauensum from the late Lower Miocene of Fiji is 
probably the nearest described species, but it has smaller, especially shorter cells, 
and narrower sporangia. 



Archaeolithothamnium sarawakense n. sp. 

(PI. 2, fig. 5) 

Diagnosis. Thallus crustose, well developed hypothallus and perithallus. 
Cells of hypothallus 16-23 /^ X 9-^4 /^- Cells of perithallus 7-14 /i X 5-1 1 fi, 
tissue fairly regular. Sporangia regularly arranged, diameter 31-48 //, height 
38-54 M- 

HoLOTYPE. Slide V. 51762. 

Sample number and locality. S-7063, south-west end of Batu Asi limestone. 
Upper Baram River, Sarawak, Borneo. 

Age. Paleocene. 

Description. Crustose. Hypothallus well developed, composed of curved 
threads of cells. Cells measure 16-23 /^ ^ong by 9-14 /i wide. Perithallic tissue 
fairly regular. Cells 7-14 /i X 5-11 M- Sporangia subspherical, in regular rows, 
lenses, or short layers (appearing as short rows in vertical sections). Size 38-54 ju, 
high, diameter 31-48 /i. 

Remarks. The dimensions of the cells and the size and shape of the sporangia 
are the same as those of A. oulianovi Pfender. However, the present species has a 
fairly regular perithallic tissue, a well developed hypothallus and the sporangia 
regularly arranged in layers or lenses; while A . oulianovi is characterized by having 
a very irregular perithallic tissue, sporangia irregularly scattered throughout the 
tissue, and a poorly developed hypothallus. It resembles the late Cretaceous 
A. brevium Lemoine but has somewhat smaller cells and a more strongly developed 
hypothallus. 



TERTIARY RED ALGAE FROM BORNEO 265 

Archaeolithothamnium saipanense Johnson 

(PL 2, fig. I) 

1957 Archaeolithothamnium saipanense Johnson : 220, pi. 38, figs. 1-4, 6. 

Description. Thallus develops a thick crust with protuberances. Hypothallus 
thin, commonly consisting of only a few curved threads of cells. Cells measure 
8-17 /t X 7-14 //. Perithallic tissue regular, with regularly spaced horizontal and 
vertical cell walls. Cells 8-13 /< X 6-13 //. Sporangia long elliptical, commonly 
closely packed in regular rows, and a single thallus may bear a number of successive 
layers. Sporangia measure 75-120 /t high and 38-55 /i wide. 

Remarks. Essentially the same as material described by the author from the 
Upper Eocene of Saipan. This species is very similar to Archaeolithothamnium 
sociahile Lemoine. The cells are essentially the same size. The main differences 
are in growth habit and sporangia. A. sociabile develops thin crusts. Normally a 
fertile crust bears only a single layer of sporangia and these are rather widely spaced. 
A. saipanense develops thick crusts with rounded protuberances. Fertile crusts 
may bear numerous successive layers of tightly packed sporangia. The sporangia 
are usually longer and narrower than those of A . sociabile. 

Sample number and locality. S-10021, S-10149, S-10153 ; all from the north 
face of the Melinau Gorge, Sarawak. 

Age. Upper Eocene (Tb) to Middle Oligocene (Td). S-10021 (Tb), S-10149 (Tc), 
S-10153 (Td). 



Archaeolithothamnium sociabile Lemoine 
(PI. 2, fig. 3) 

1939 Archaeolithothamnium sociabile Lemoine : 53-54, text-figs. 16-17. 

1961 Archaeolithothamnium cf. A. sociabile Lemoine ; Johnson : 919, pi. 267, figs. 3, 4. 

Description. Crusts, commonly thin, may be superimposed. Hypothallus 
thin and may be difficult to recognize as commonly composed of only two or three 
horizontal or very slightly curved cell threads. Cells measure 8-24 /i X 7-14 /*, 
commonly 8-16 /i x 7-11 /i. Perithallic tissue regular with horizontal partitions more 
conspicuous than the vertical. Cells 7-22 fi x 7-12 //, commonly 11-17 [j, x 7-12 /i. 
Sporangia in regular rows (as seen in vertical section) , commonly only a single row 
to a thallus. Sporangia 80-100 [i high and 35-65 pt wide. 

Remarks. Numerous specimens observed at one locality which very closely 
fit Lemoine's descriptions of the type material from the Upper Eocene and Oligocene 
of Algeria. 

Sample number and locality. S-10153, north face of Melinau Gorge, north-east 
Sarawak. 

Age. Oligocene (Td). 



266 TERTIARY RED ALGAE FROM BORNEO 

Lithothamnium aggregatum Lemoine 

1939 Lithothamnium aggregatum, Lemoine : 66-67, pl- !■ fig- 12; pi. 3, figs. 3, 4, text-fig. 27. 

Description. Thin crusts, commonly less than 600 fi thick, which may grow 
superimposed or interstratified with other species or other organisms. Hypothallus 
slightly developed, commonly around 100-125 {i thick, of curved cell threads. 
Cells measure 15-22 /* long and 9-13 /i wide. Perithallic tissue fairly regular, 
300-600 /I thick, with cells 14-27 [i x 8-13 //. No conceptacles observed. 

Remarks. Several infertile specimens observed in the Borneo collection which 
fit Lemoine's description of the species from the Oligocene of Algeria. 

Sample number and locality. S-10160, Melinau Gorge, north-east Sarawak. 

Age. Oligocene (Td). 



Lithothamnium cantabricum Lemoine 
(PI. 2, fig. 2) 

1934 Lithothamnium cantabricum Lemoine in Lemoine & Mengaud : 175, text-fig. i. 
1961 Lithothamnium cantabricum Lemoine ; Segonzac : 442. 

Description. Thallus develops as an undulating crust 0-25 to 0-4 mm. thick. 
The hypothallus is well developed (0-14 to 0-17 mm. thick), with cells 13-24 /i long 
and 9-11 ju, thick. Perithallus o-ii to 0-17 mm. thick with cells 7-11 /i x 8-11 /<. 

Remarks. Represented by a single specimen in the Borneo collection. It 
differs from the holotype only in having a better developed hypothallus. 

Sample number and locality. S-7063, south-west end of Batu Asi limestone. 
Upper Baram River, Sarawak. 

Age. Paleocene. 



Lithothamnium. cf. caucasicum Maslov 
(PI. 2, fig. 4) 

1956 Lithothamnium caucasicum Maslov : 116-117, pi. 33, fig. 2, text-fig. 46. 

Description. Thallus develops short stubby branches. The branches consist 
mainly of a medullary area of arched " layers " of cells and a narrow marginal area. 
The central portion is composed of threads of cells 29-56 fi x 12-17 /^- ^^ 'the lower 
part of the branch, this central tissue is irregular with the cell threads branching 
frequently and the cross partitions between the cells irregularly spaced. Higher up 
the tissue becomes regular with cross partitions strong and regularly spaced, giving 
the suggestion of " layers " of cells. There are also suggestions of growth zones. 

The marginal tissue is thin and fairly regular. Cells 14-17 fi X 11-14 fi. No 
conceptacles present. 



TERTIARY RED ALGAE FROM BORNEO 267 

Remarks. The cell dimensions and frequent branching of the cell threads in the 
lower central area agree with Maslov's species from the Danian of the Caucasus 
region. The upper portion (not described or illustrated by Maslov) may be more 
regular. 

Sample number and locality. S-6177, Upper Baram River, Sarawak. 

Age. Paleocene. 



Mesophyllum curtum Lemoine 

(PI- 3. fig. 3) 
1939 Mesophyllum curtum Lemoine : 92, text-fig. 61. 

Description. Long slender branches. Tissue somewhat irregular, the vertical 
cell threads commonly as pronounced as the curved horizontal " layers ". Numerous 
growth zones but their boundaries not pronounced. Cells unusually small, 7-9 /j, x 
8-10 ju. Conceptacles 325-343 /t in diameter, 140-149 ju, high. 

Remarks. This form has unusually small cells. In character of tissue and size 
of cells and conceptacles, it fits almost exactly Lemoine's type material. It differs 
only in having longer and more narrow branches. 

Sample number and locality. S-10177, Melinau Gorge, Sarawak. 

Age. Lower Miocene (Lower Te). 



Mesophyllum cf. pfenderae (Lemoine) 
(PI. 3, figs. I, 2) 

1928 Lithophyllum pfenderae Lemoine : 100, text-fig. 14. 

1939 Mesophyllum pfenderae (Lemoine) Lemoine : 87. 

1961 Mesophyllum pfenderae (Lemoine) ; Segonzac : 438-439, text-fig. 4. 

Description. The plants develop short sinuous branches i-o to 1-3 mm. thick. 
Medullary hypothallus forms most of the branch. It is composed of arched layers 
of cells arranged in pronounced growth zones, 5 to 9 cell layers to a zone. Cells 
rectangular, 28-40 /x x 9-15 yW. Marginal perithallus commonly formed of 3 to 6 
layers of cells which measure 13-40 n x 9-14 /^. Conceptacle 352 [i x 125 [i. 

Remarks. The Borneo material closely resembles the descriptions given by 
Lemoine and by Segonzac. It differs in having somewhat longer hypothallic cells. 
The branches may be more sinuous. Unfortunately, neither Lemoine nor Segonzac 
illustrates the branches or gives any descriptions of them, beyond diameter and length 
measurements. 

Sample number and locality. S-6175, Upper Baram River, Sarawak. 

Age. Paleocene. 



268 TERTIARY RED ALGAE FROM BORNEO 

Mesophyllum vaughanii (Howe) Lemoine 
(PL 4, fig. 3 ; PI. 5- fig. 5) 

1918 Lithothamnium vaughanii Howe : 6-7, pis. 7, 8. 

1939 Mesophyllum vaughanii (Howe) Lemoine : 89, pL i, figs. 2, 8, 11, 15. 

1962 Mesophyllum vaughanii (Howe) ; Johnson : 157, pL 3, figs, i, 2. 

Description. The plant starts as a basal crust from which develop irregular 
protuberances or long slender branches. Basal hypothallus of curved rows of cells. 
The branches are formed of numerous, irregularly disposed growth zones. These 
consist of " layers of cells ", commonly quite regularly arranged. Cells of central 
part of growth zones of branches slightly longer than those of basal hypothallus. 
Conceptacles large, numerous, and multiple apertured. 

Table I. — Dimensional data of Mesophyllum vaughanii (in mu) 



Slide 


Hypothallic Cells 


Perithallic Cells 


Conceptacles 


S-ioio9a 




11-21 X 10-15? 


475-528 X220 


S-ioi48b 


13-22x8-13 


13-20x8-13 


364-510 X 144-176 


S-ioo32a 




14-21 x6-ii 




S-ioiooa 


27-32 X8-19 


7-12 X9-12 




S-ioo32a 




19-26x8-14 




S-i 0087b 




i2-ig X8-13 


448-589 X 167-220 


Range 


13-32x8-19 


11-21(26) X (6)8-15 


364-589 X 167-220 



Remarks. This is a widely distributed Upper Eocene-Lower Oligocene species. 
It is represented abundantly in the Borneo collection. 

Sample number and locality. S-10032 (Tb), S-10087 (Tb or Tc), S-ioioo (Tc), 
S-10109 (Tc), S-ioioo (Tc), S-10148 (Tc), S-10153 (Td). Melinau Gorge, north-east 
Sarawak. 

Age. Upper Eocene (Tb) to Middle Oligocene (Td). 

Mesophyllum vaughanii (Howe) var. sarawakense nov. 
(PL 4, figs. I, 6) 

Diagnosis. Strongly branching with thick medullary hypothallus and thin 
marginal perithallus. Medullary tissue regular with cells 15-40 /i x 7-26 fi. Peri- 
thallic cells 9-18 /I X 7-12 //. Suggestions of marginal conceptacles. 

Holotype. Slide V.51772. 

Sample number and locality. S-10153 and 10154, north face of Melinau 
Gorge, north-east Sarawak. 

Age. Middle Oligocene (Td). 

Description. Thallus starts as a rather thin crust from which develop long 
medium to thick branches. Branches consist of a strongly developed medullary 
hypothallus surrounded by a thin marginal perithallus. Growth zones strongly 
developed in some specimens, not so prominent in others. Branches I-6-3-8 mm. 



TERTIARY RED ALGAE FROM BORNEO 269 

thick and more than 6-o mm. long. Growth zones 8-12 layers thick. Medullary 
cells 15-40 fi X 7-26 //., commonly 20-28 /i X 10-17 /^- Cells in regular " layers " 
with thick horizontal partitions and thin vertical partitions. Marginal perithallus 
thin, cells commonly 9-18 [i x 7-12 fi. Several specimens show suggestions of 
marginal conceptacles with diameters up to about 600 fi. However, they are badly 
overgrown and precise measurements cannot be made. 

Remarks. This species closely resembles the typical Mesophyllum vaughanii 
(Howe) Lemoine in general appearance and growth habits. It differs slightly but 
consistently in several respects: the medullary cells are longer but narrower, the 
medullary tissue is more regular without the frequent development of secondary 
hypothallia so characteristic of M. vaughanii, and while there are suggestions of 
alternating layers of longer and shorter cells in some areas or all of the medullary 
tissue, this is not so common as in M. vaughanii and there is less difference in the 
actual size of the cells in alternate layers. 

This form occurs abundantly in samples S-10153 and 10154. 

Lithophyllum besalotos Johnson 

1962 Lithophyllum besalotos Johnson : 159, pi. 4, figs. 4, 5. 

Description. Plant forms thin irregular crusts which may grow superimposed. 
Hypothallus thin but easily recognizable, of irregularly curved cell threads. Cells 
measure 14-25 fi x 7-1 1 i^- Perithallic cells wider than high with thick horizontal 
cell walls. Cells arranged in layers like bricks. Cells measure 9-14 /^ X 12-22 /(. 
No conceptacles observed. 

Remarks. Represented by only a few poorly preserved specimens. 

Sample number and locality. S-10162, Melinau Gorge, Sarawak. 

Age. Middle Oligocene (Td). 

Lithophyllum capederi Lemoine 

(PI. 5, fig- 6) 

1900 Lithothamnium tenue Capeder : 180, pi. 6, fig. 15. 
1900 Lithothamnium dentatum Capeder : 178, pi. 6, fig. 7. 
1926 Lithophyllum, capederi Lemoine : 11, text-fig. 11. 

Description. Thallus forms a thin crust. Hypothallus coaxial with exception- 
ally thick concentric partitions. Hypothallic cells 13-19 /^ X 8-11 fi. Perithallic 
tissue regular, composed of small, square or rectangular cells 6-10 /i X 4-9 11. 

Remarks. This species is characterized by a coaxial hypothallus with unusually 
thick cell walls and very small perithallic cells. In cell measurements it closely 
resembles Lithophyllum johnsoni Ishijima. 

Sample number and locality. S-7080, Belukan River, Sarawak. 

Age. Lower Miocene (Te). 



270 TERTIARY RED ALGAE FROM BORNEO 

Lithophyllum densum Lemoine 
(PI. 5, fig- 8) 

1934 Lithophyllum densum Lemoine : 282, text-fig. 14. 

1956 Lithophyllum, aff. L. densum, Lemoine ; Maslov : 117, pi. 34, figs. 1-4. 

Description. Long straight cylindrical branches with diameters ranging from 
0-9 to i-i mm. A medullary hypothallus with gently arched layers of cells attains 
a diameter of 0-7 to o-8 mm., with cells 25-40 /i long and 8-15 jj, wide. The marginal 
perithallus is 88-180 /i thick, with cells 13-16 /i long and 7-12 /i wide. 

"^ xARKS. This species is characterized by the long straight branches, thick 
medullary hypothallus with gently arched layers, and the long narrow hypothaUic 
cells. The cells of the Borneo specimens are appreciably longer than those in the 
holotype of Mid-Eocene age, and slightly longer than Maslov's early Paleocene 
material. The other dimensions are the same. 

Sample number and locality. S-6177, Upper Baram River, Sarawak. 

Age. Paleocene. 

Lithophyllum dubium Lemoine 

1934 Lithophyllum, duhium Lemoine : 282, text-fig. 13. 

1961 Lithophyllum dubium Lemoine ; Segonzac : 443, text-fig. 9. 

Description. Thallus thin, crustose. Several may grow superimposed. Hypo- 
thallus about 200 fi thick with cells 17-25 /a x 7-12 fi. Perithallus regular, up to 
300 II thick, with cells 15-17 n X 11-15 /.i. 

Remarks. Represented by several infertile specimens. 

Sample number and locality. S-7073, Upper Baram River, Sarawak. 
Age. Paleocene. 

Lithophyllum cf. obliquum Lemoine 
(PI- 4, fig- 5) 

1930 Lithophyllum obliquum, Lemoine : 266, text-figs. 1,2. 

1939 Lithophyllum cf. L. obliquum Lemoine ; Lemoine : 97, 98, text-fig. 64. 

Description. Crustose, o-4-o-55 mm. thick. Hypothallus 130-155 n thick, 
poorly coaxial, " layers " only slightly curved, almost oblique with thick walls. 
Cells 20-25 /* X 6-14 II. Perithallus 350-450 fi thick with well defined " layers " 
of cells. Cells measure 8-12 fi x 8-14 ft. 

Remarks. The general structure suggests L. obliquum Lemoine and the cells 
are about the same size. The main difference is a greater development of perithallic 
tissue. It is similar to Lithophyllum cf. obliquum Lemoine from Algeria. 

Sample number and locality. S-7081, Belukan River, Sarawak. 

Age. Lower Miocene (Te). 



TERTIARY RED ALGAE FROM BORNEO 271 

Lithophyllum cf. ovatum (Capeder) 

1900 Lithothamnium ovatum Capeder : 177, pi. 6, figs. $a, b. 

1926 Lithophyllum ovatum (Capeder) Lemoine : 245-246, text-fig. 3. 

1932 Lithophyllum ovatum (Capeder) ; Airoldi : 70, pi. lo. 

1957 Lithophyllum ovatum (Capeder) ; Johnson : 228, pi. 45, figs. 4, 8. 

Description. Thallus crustose with a well developed hypothallus and perithallus. 
Hypothallus 90-250 /i thick, poorly coaxial. Cells 11-22 /< x 12-16 fi. Perithallus 
100-250 /i thick with cells in slightly irregular " layers ". Cells measure 7-17 fi x 
7-1 1 /<. No conceptacles observed. 

Remarks. This form strongly suggests L. ovatum but the material avails, for 
study is too limited to be certain. 

Sample number and locality. S-ioioo, Melinau Gorge, north-east Sarawak. 

Age. Lower Oligocene (To). 

Lithophyllum quadrangulum Lemoine 
(PI. 4. fig- 4) 

1934 Lithophyllum quadrangulum. Lemoine : 279, text-fig. 10. 

1934 Lithophyllum, quadrangulum Lemoine ; Lemoine & Mengaud : 178, text-fig. 4. 

Description. Plant forms a very thin crust (175-220 /i thick), composed entirely 
of hypothallic tissue. Hypothallus coaxial but the curved layers of cells form only 
gentle arcs and they are almost vertical. Cells large, measuring 28-34 1^ X 13-17 r- 

Remarks. This species has very characteristic features : (i) a thin crust con- 
sisting entirely or almost entirely of hypothallus ; (2) the unusual structure of the 
hypothallus ; and (3) the large cells. It has a long time range (Mid-Eocene to Mid- 
Miocene), and apparently became widely distributed geographically. 

Sample number and locality. S-10160, Melinau Gorge, north-east Sarawak. 

Age. Middle Oligocene (Td). 

Lithoporella melobesioides (Foslie) 
(PL 2, fig. 6) 

1904 Mastophora [Lithoporella) melobesioides Foslie ; Weber van Bosse & Foslie : J^-TJ, text- 
figs. 30-32. 
1939 Melobesia [Lithoporella) melobesioides Foslie; Lemoine; 108-110, text-figs. 78, 79. 
1943 Lithoporella melobesioides (Foslie) ; Lignac-Grutterink : 292-293, pi. 2, fig. 8. 

1949 Lithoporella [Melobesia) melobesioides (Foslie) Johnson & Ferris : 196-197, pi. 37, 

figs. 4, 5 ; pi. 39, fig. 2. 

1950 Lithoporella melobesioides (Foslie) ; Johnson & Ferris : 18, pi. 8, fig. A. 

1957 Lithoporella melobesioides (Foslie) ; Johnson : 234, pi. 37, fig. 5 ; pi. 43, figs, i, 2 ; pi. 
49. fig- 4 ; pl- 56, fig- 6. 
Description. Thallus consists of a single layer of large cells except around the 
conceptacles. Cells elongated vertically and commonly slightly obliquely. Cells 
range greatly in size even in a single slice across a thallus. Range of cell size of 
17 specimens measured was 26-82 /i high and 12-35 /* wide. 

GEOL. 11,6 26 



272 



TERTIARY RED ALGAE FROM BORNEO 



Remarks. This is probabl}' the most common species of coralhne algae observed 
in the Borneo collections. Fragments occur on many slides. The cell dimensions 
all fit within those of the highly variable, widespread, long ranged L. melobesioides. 

Sample number and locality. S-7063, south-west end of Batu Asi limestone. 
Upper Baram River, Sarawak. 

Age. Paleocene. 



Table 2, 
Cell 

Slide 
5-70636 
S-7o63g 
S-6i75a 
S-7o63h 
5-7063111 
5-70776 
S-6i75a 
BM. 13 
S-10153C 
5-ioi76b 
5-71 I li 



— Lithoporella melobesioides (Foslie) 
Size of Typical Specimens (in mu) 

Cell dimensions Age 

Paleocene 



48-75 X 21-26 
49-70 X 29-35 
45-82 X 24-33 
66-70 X 16-19 
56-73 X 30-48 
26-39,x 14-18 
41-53x17-24 
30-48 X 14-22 

30-51 X 14-29 
28-43 X 22-30 

27-40x17-21 



Paleocene 

Paleocene 

Paleocene 

Paleocene 

Paleocene 

Paleocene 

Paleocene 

Middle Oligocene 

Lower Miocene 

Lower Miocene 



Lithoporella antiquitas Johnson 
(PI. 6, fig. 3) 

1961 Lithoporella antiquitas Johnson : 937, pi. 276, figs, i, 2. 

Description. Thallus very small, encrusting, consisting of a single layer of 
vertically elongated cells, 44-53 n long and 14-17 fi wide. Conceptacle small, 
341 fi in diameter and 128 /.i high. 

Remarks. Closely resembles the holotype from the Miocene of Eniwetok, 
Marshall Islands, except for a slightly larger conceptacle. 

Sample number and locality. S-7081, Belukan River, Sarawak. 
Age. Lower Miocene (Te). 

Lithoporella cf. minus Johnson 
(PI. 3, fig. 5) 

1964 Lithoporella minus Johnson : Cio, pi. 2, fig. 6. 

Description. Thallus tiny, irregular crustose, consisting of a single layer of 
cells except around the conceptacle. Cells 8-20 fi high, and 11-18 /t wide. Con- 
ceptacle 373 fi X 200 [I, with a single large pore. 

Remarks. Only one specimen observed in the Paleocene collection. It closely 
resembles the holotype from the Eocene of Ishigaki Ryuk}^ Islands, except for 
sUghtly shorter cells. 

Sample number and locality. S-7077, Upper Baram River, Sarawak. 

Age. Paleocene. 



TERTIARY RED ALGAE FROM BORNEO 273 

Melobesia cf. cuboides Johnson 

1957 Melobesia ? cuboides Johnson : 234, pi. 43, figs. 6, 7. 
1962 Melobesia cuboides Johnson ; Johnson : 164, pi. 5, fig. 6. 

Description. Thallus monostromatic. Cells cubic or horizontally elongated 
with thick walls. Cell size 15-34 fi X 12-21 fi. 

Remarks. Similar to material previously described by Johnson (1962) from Batu 
Gading, Sarawak. 

Sample number and locality. S-6623, south-west end of Melinau limestone, 
S-10176, Melinau Gorge, north-east Sarawak. 

Age. Lower Miocene (Lower Te). 

Dermatolithon saipanense Johnson 
(PI. 6, fig. 4) 

1957 Dermatolithon saipanense Johnson : 235, pi. 57, figs. 4, 6. 

Description. Thallus forms a thin crust. Hypothallus of one or two layers of 
elongated cells 32-46 fi high and 14-21 /* wide. Perithallus of square or vertically 
elongated cells, 30-33 /* long and 24-32 /i wide. 

Remarks. The one Borneo specimen observed has cells within the size range of 
this species, although the average cell size is a little smaller than that of the Saipan 
material. 

Sample number and locality. S-7111, Belukan River, Sarawak. 

Age. Lower Miocene (Lower Te). 

THAUMATOPORELLA Pia 1927 

Thaumatoporella parvovesiculifera (Raineri) 

(PL 6, fig. 6) 

1922 Gyroporella parvovesiculifera Raineri : 83, pi. 13, figs. 17, 18. 

1927 Thaumatoporella parvovesiculifera (Raineri) Pia : 69. 

1938 Thaumatoporella parvovesiculifera (Raineri) ; Pia : 491, pi. i, figs. 1-5 ; pi. 2, figs. 6-14. 

1956 Lithoporella melobesioides (Foslie) ; Elliott : 327, pi. 2, figs. 8, 9. 

1957 Polygonella incrustata Elliott : 230, pi. i, figs. 11, 12. 
1957 Lithoporella elliotti Emberger : 625, pi. 32, figs. 1-4. 

1959 Thaumatoporella cf. parvovesiculifera (Raineri) ; Gasche, pi. i, fig. 3. 

1959 Thaumatoporella parvovesiculifera (Raineri) ; Sartoni & Crescenti : 129, pi. 2, figs. 1-5. 

i960 Thaumatoporella parvovesiculifera (Raineri) ; Radoicic : 133, pis. i, 2. 

Description. Thallus consists of a single layer of long cells, polygonal in cross- 
section. Cells 34-91 /( long, and 16-22 n wide. 

Remarks. Grows encrusting a shell or other hard object. Closely resembles 
Elliott's (1957) late Jurassic-early Cretaceous material except in having slightly 
smaller cells (34-91 /i X 16-22 [i instead of 65-104 /^ x 26-33 [') arid more undulating 
growth habit. Only a single specimen observed. 

GEOL. II, 6 26§ 



274 TERTIARY RED ALGAE FROM BORNEO 

Sample number and locality. S-7063, south-west end of Batu Asi limestone, 
Upper Baram River, Sarawak. 
Age. Paleocene. 

Subfamily CORALLINOIDEAE (Articulated Corallines) 

Genus CORALLINA Linnaeus 1758 

Corallina sp. A. 

Description. Segments small, o-8-i-3 mm. long and o-i6-o-25 mm. wide, 14 to 18 
tiers of cells to a segment. Cells at centre of tiers 47-64 /t long and 6-12 n wide. 
Marginal cells 22-35 /* lorig ^^^ 6-14 fi wide. 

Remarks. Only a few worn segments of Corallina were observed in the Paleocene 
collection. Data from the two best specimens are given above. They closely 
resemble Corallina sp. /. (Segonzac 1961 : 444). 

Sample number and locality. S-7063, south-west end of Batu Asi limestone, 
Upper Baram River, Sarawak. 

Age. Paleocene. 

Corallina cf. abundans Lemoine 

(PI. 3, figs. 6, 7) 

1934 Corallina abundans Lemoine : 284-285, text-fig. 16. 

Description. Segments o-i4-o-22 mm. wide. Cells of medullary h5/pothallus at 
middle, 52-64 /< long and 7-12 /t wide. Marginal cells 13-14 /i X 8-11 {i wide. 

A fragment of Corallina with a conceptacle chamber occurs on the same slide and 
probably represents the same species. The conceptacle space is rounded conical, 
218 fi wide near base and 356 /i high. 

Remarks. Represented by only a few frayed fragments from a single locality. 
Their cells, both medullary and marginal, agree in length with Lemoine's species but 
are much narrower (hypothallus 7-12 fi versus 8-20 fi, perithallus 8-11 jjl versus 
15-25 fi). 

Sample number and locality. S-6610, south-west end of Melinau limestone, 
Sarawak. 

Age. Lower Oligocene (Tc). 

Corallina neuschelorum Johnson 

(PI. 5- fig. 4) 
1957 Corallina neuschelorum Johnson : 239, pi. 37, fig. 3 ; pi. 50, figs. 1-4. 

Description. Segments flattened, mainly hypothallic tissue, with cells in centre 
of layers 48-76 ju, long and 9-15 ju, wide. Marginal cells 19-27 /i x 8-16 /i. 

Remarks. The cell dimensions closely fit those of the Saipan species, although the 
marginal cells are somewhat longer (19-27 /< versus 15-21 jti). 

Sample number and locality. S-7111, Belukan River, Sarawak. 

Age. Lower Miocene (Lower Te). 



TERTIARY RED ALGAE FROM BORNEO 275 

Corallina cf. prisca Johnson 
1957 Corallina prisca Johnson : 239, pi. 37, fig. 4 ; pi. 44, figs, i, 2, 7-11. 

Description. Pieces of segments about 0-35 mm. wide. Cells at centre of 
medullary tissue 63-88 fi long and 7-8 ft wide. Node between two segments 201 ju 
long. 

Remarks. The dimensional data for these fragments is within the range of 
Corallina prisca from the Upper Eocene of Saipan. 

Sample number and locality. S-10025 (Tb) and S-10153 (Td), both from 
north face of Melinau Gorge, Sarawak. 

Age. Upper Eocene (Tb) and Middle Oligocene (Td). 

Jania miocenica Johnson 
(PI- 5, fig. 3) 

1961 Jania miocenica Johnson : 938-939, pi. 278, figs. 6-8. 
Description. Slender segments with dimensional data tabulated below. 

Table 3. — Jania miocenica (Dimensions in mu) 

Hypothallic Cdls Perithallic Cells Age 

Length Width 

33-39 13-17 

35-46 12-18 

36-50 14-20 

33-40 14-22 

Remarks. In general appearance and dimensional data, the Borneo specimens 
agree exactly with those described by Johnson (1961) from the Lower Miocene of 
Eniwetok. 

Sample number and locality. S-6921 (Tc), mouth of Tukuruk River, MeHnau, 
Sarawak. S-7111 (Te), Belukan River, Sarawak. 

Age. Lower Oligocene (Tc) and Lower Miocene (Lower Te). 

Jania cf. nummulitica Lemoine 

(PI. 6, fig. 5) 

1934 Jania nummulitica Lemoine : 285. 

Description. Piece of a segment 2-12 mm. long and 0.33 mm. wide. It contains 
35 layers of cells, with approximately 18 cells in a row as cut by the section studied. 
The cells at the centre of the rows measure 63-76 fi high and 11-18 n wide. The 
marginal cells are 25-30 /i high and 16-21 [i wide. 

Remarks. Represented by a couple of fragments. The largest is described 
above. The cell dimensions are close to those of /. nummulitica Lemoine from the 
Upper Eocene of Hungary. However, the cells of the Borneo specimens are slightly 
shorter and wider than the holotype. 



Slide 


Number 


number 


cells in row 


S-692id 


19 


S-692id 


17-19 


S-692id 


27-32 


S-yiiid 


22 



Length 


Width 




15-26 


14-24 


Lower Oligocene 


14-30 


9-21 


Lower Oligocene 


13-28 


16-21 


Lower Oligocene 


11-21 


11-18 


Lower Miocene 



276 TERTIARY RED ALGAE FROM BORNEO 

Sample number and locality. S-6921, mouth of Tukuruk River, Melinau, 
Sarawak. 

Age. Lower Oligocene (Tc). 

Jania vetus Johnson 
(PL 4, fig. 2) 

1957 Jania vetus Johnson : 237, pi. 52, fig. 2. 
1 961 Jania vetus Johnson ; Johnson : 939. 

Description. Long, sometimes branching segments 0-26 to 0-48 mm. wide. 
Medullary cells from near centre measure 50-77 /^ long and 15-26 [i wide. Marginal 
cells 13-16 [i X 26-46 fi. 

Remarks. The cell sizes are within the ranges of the species described from the 
Miocene of Saipan. 

Sample number and locality. S-7111, Belukan River, Sarawak. 

Age. Lower Miocene (Lower Te). 

Amphiroa cf. fortis Johnson 

(PI- 6, fig. 7) 

1961 Amphiroa fortis Johnson : 939, pi. 277, figs. 8, 9. 

Description. Segments probably long, with diameters ranging from 0-7 to 
0-9 mm., commonly o-8 to 0-87 mm. Medullary hypothallus of layers of cells. The 
cells at the centre measure 42-79 jj, long and 9-30 /i wide. There is a suggestion of 
alternating layers of long and short cells, but the difference in length is slight, 
62-79 /^ versus 49-64 [i. Marginal perithallic cells 15-29 /^ X 12-26 [i. 

Remarks. This form is very close to Amphiroa fortis Johnson from the late 
Eocene of Eniwetok. The Borneo specimens have cells about the same length but 
wider, 9-30 [i versus 6-1 1 /t in the hypothallus, and 12-22 [i versus 7-11 /i in the 
perithallus. 

Sample number and locality. S-7073, Upper Baram River, Sarawak. 

Age. Paleocene. » 

Amphiroa sp. 

Description. Segments around 0-3 mm. wide. Medullary tissue shows an 
alternation of layers of long and short cells, formula i-long, i-short. Cells at centre 
of medullary area measure: long 42-56 fi x 7-8 /i, short 34-40 /^ x 7-8 /<. 

Remarks. Represented by a few fragments. 

Sample number and locality. S-10055, Melinau Gorge, Sarawak. 

Age. Upper Eocene (Tb). 



TERTIARY RED ALGAE FROM BORNEO 277 

Subterraniphyllum thomasi Elliott 

(PL 6, figs. I, 2) 
1957b Subterraniphyllum thomasi Elliott : 73-74, pi. 13, figs. 1-9. 

Description. Segments composed of wide medullary hypothallus and very 
narrow marginal perithallus. Hypothallus composed of slightly curved layers 
of large irregular cells. Cell layers shaped much the same as in Amphiroa with 
flattened top and sharply inclined margins. Cells irregular to almost regular in 
vertical section ; very irregular, rounded to polygonal in horizontal section. Cells 
85-118 /I long and 58-99 fi wide. Marginal perithallus thin, cells rectangular, 
15-26 fi X 7-21 ju,. 

Remarks. In sections the medullary hypothallus appears light while the peri- 
thallic tissue is unusually dark, practically black. The Borneo material consists of 
fragments of segments. They fit Elliott's description of the type material from 
Iraq except that the perithallic cells are slightly larger. 

Sample number and locality. S-6921, mouth of Tukuruk River, Melinau, 
Sarawak. 

Age. Lower Oligocene (Tc). 

Genus DISTICHOPLAX Pia 1934 

Distichoplax biserialis (Dietrich) 

(PI. 5, fig. 7) 

1918 Lithothamniiim ? sp., Trauth : 220, pi. 11, figs. 2, 3. 

1918 Lithothamnium nummuliticum Trauth : 219, pi. 11, fig. i. 

1927 Lithothamnium hiseriale Dietrich : 461, pi. 11, fig. i. 

1930 Lithoporella Pia : 133. 

1934 Distichoplax biserialis (Dietrich) Pia : 15, text-figs. 5-8. 

1956b Distichoplax biserialis (Dietrich) ; Elliott : 332, pi. 11, fig. i. 

1958 Distichoplax biserialis (Dietrich) ; Lemoine : 2145. 

i960 Distichoplax biserialis (Dietrich) ; Elliott : 226, pi. 8, figs. 2, 3. 

1961 Distichoplax biserialis (Dietrich) ; Segonzac : 446, pi. 13, figs. 3, 4. 

1963 Distichoplax biserialis (Dietrich) ; Keij : 153-160, pi. i. 

Description. The thallus develops as small undulating plaque or sheet. They 
may occur singly or a number may grow in close association. Commonly each one 
will be oriented somewhat differently. Sections cutting the thallus at different 
angles look very differently. The most characteristic is a transverse section cut 
parallel to the length of the cells. This gives the " fishbone fossil " with two rows of 
obliquely elongated cells coming together at a wide angle (PL 5, fig. 7). In the 
specimen illustrated, the cells have lengths of 76-81 [i and widths of 30-41 jx,. 

Remarks. This is a very widespread Paleocene and early Eocene fossil, with its 
greater development in the Paleocene. 

Sample number and locality. BM13, Batu Asi limestone. Upper Baram area, 
Sarawak. 

Age. Paleocene. 



278 TERTIARY RED ALGAE FROM BORNEO 

Family SOLENOPORACEAE 

Genus SOLENOMERIS Douville 

" Solenomeris " sp. 

(PI. 5, fig. I) 

Description. Thallus forms rounded masses from which branches or irregular 
protuberances may develop. The fragments observed in the Borneo collection 
consisted of perithallic tissue. This is composed of threads of coarse semirectangular 
cells, regularly arranged. The walls of the cell threads are quite thick. The cross 
partitions between the cells are thinner. They develop at approximately the same 
levels in adjoining threads, giving the tissue an articulated or layered appearance in 
vertical sections. Cells measure 21-33 ju. high and 41-65 /i wide. 

Remarks. This is one of the characteristic Paleocene fossils. It occurs in beds 
of about Danian age from Morocco to the East Indies and in Guatemala and Cuba. 
[Since this paper was submitted, Professor Johnson has pointed out that Solenomeris 
may be a hydrozoan, in which case he thinks that the present species should be referred 
to Parachaetetes. Ed.] 

Sample number and locality. B.M. 47, south-west end of Batu Asi limestone, 
Upper Baram River, Sarawak. 

Age. Paleocene. 



Family SIPHONOCLADACEAE Schmitz 1879 
Genus PYCNOPORIDIUM Yabe & Toyama 1928 

Pycnoporidium sinuosum Johnson & Konishi 
(PI. 5. fig. 2) 

i960 Pycnoporidium sinuosum Johnson & Konishi : iioo-iioi, pi. 134, figs. 1-4. 

Description. Thallus consists of an irregular mass of loosely interwoven, coarse, 
sinuous, filaments. These filaments are 41-58 pi in diameter. They are partitioned 
by cross walls (septa) at irregular intervals (47 /i to 72 jj). Second branching occurs 
sparsely at irregular intervals. The filaments have thick walls with occasional 
uncalcified chambers occupying the spaces within the tubes between the septa. 

Remarks. This species was originally described from the late Cretaceous of 
Guatemala. It differs from the only previously described Paleocene species from 
Iraq, P. levantian Johnson, in having slightly smaller tubes and more closely spaced 
septa. Represented by only a few specimens in the Borneo collection. 

Sample number and locality. S-7077, Upper Baram River, Sarawak. 

Age. Paleocene. 



TERTIARY RED ALGAE FROM BORNEO 279 

V. REFERENCES 

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Butll. Inst, catal. Hist. nat. L^rida (2) 8, 5-6 : 92-107, 20 figs. 

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28o TERTIARY RED ALGAE FROM BORNEO 

Lemoine, Mme Paul 1939. Les algues calcaires fossiles de L'Algdrie. Bull. Cartegeol. Alger. 

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Lemoine, Mme Paul & Mengaud, M. L. 1934. Algues calcaires de I'Eocene de la Province de 

Santander (Espagne). Bull., Soc. Hist. nat. Toulouse, 46 : 171-180, 6 figs. 
LiECHTi, P., Roe, F. W. & Haile, N. S. i960. The Geology of Sarawak, Brunei and the western 

part of North Borneo. Bull. Geol. Surv. Dept. Brit. Terr. Borneo, 3. 360 pp., 73 pis. 
Lignac-Grutterink. 1943. Some Tertiary Corallinaceae of the Malaysian Archipelago. 

Verh. geol.-mijnb. Genoot. Ned. Kolon., 's Gravenhage [Geol.) 113 : 292-293, pi. 2, fig. 8. 
Maslov, V. P. 1956. Fossil algae of the U.S.S.R. Trav. Inst. Sci., Geol. Akad. Nauk. SSSR., 

160 : 1-301. 
Miranda, F. 1935. Algas coralinaceas fosiles del terciario de San Vicente de la Barquera 

(Santander). Bol. Soc. esp. Hist, nat., Madrid, 30 : 279-287. 
Pfender, J. 1926. Sur les organismes du Nummulitique de la colline de San Salvador pres 

Camarasa Catalogne. Bol. Soc. esp. Hist, nat., Madrid, 26 : 321-330, pis. 8-15. 
Pia, J. 1927. Thallophyta. In Hirmer, M. Handbuch der Paldobotanik, 1 : 31-136. 
1930. Neue Arbeiten iiber fossile Solenoporaceae und Corallinaceae. Neues Jb. Miner., 

Stuttgart, 3 : 122. 
1934- Kalkalgen aus dem Felsen von Hricovsk6 Podhradie im Waagtal. Vesth. geol. 

List. cesl. Rep. Praha, 10 : 14-18, figs. 5-8. 
1938. Uber Thaumatoporella parvovesiculifera Rain. spec, und ihr Auftreten auf der Insel 

Naxes. Praktika Akad. Athen., 13 : 491-495, pis. i, 2. 
Radoicic, R. i960. On the little known species Thaumatoporella parvovesiculifera (Rain.) 

Bull. Serv. Geol. Geophys. Serbie, Beograd (A) 18 : 133-140, pis. i, 2. 
Rama Rao & Pia, J. 1936. Fossil algae from the uppermost Cretaceous beds (Niniyur group) 

of the Trickinopoly district S. India. Palaeont. indica, Calcutta, 21, 4 : 1-49, pis. 1—6. 
Raineri, R. 1922. Alghe siforee fossili della Libia. Atti. Soc. ital. Sci. nat., Milano, 61 : 

72-86, pi. 3. 

1923. Alghe fossili mioceniche di Cirenaica. Niiova Notarisia, Padova, 35 : 5-23, 16 figs. 

RoTHPLETZ, A. 1891. Fossile Kalkalgen aus den Familien der Codiaceen und der Corallineen. 

Z. dtsch. geol. Ges., Berlin, 43 : 295-322, pis. 15-17. 
Sartoni, S. & Crescenti, U. 1959. La zona a Palaeodasycladus mediterraneus (Pia) nel Lias 

dell'Appennino meridionale. Giorn. Geol., Bologna (2) 27: 115-139, pis. 1-3. 
ScHWAGER. 1883. Die Foraminiferen aus den Eocanablagerungen der libychen Wiiste und 

Aegyptens. Palaeontographica, Stuttgart, 30 : 79-154, pis. 24-29. 
Segonzac, G. 1961. Niveaux a' Algues dans le Thanetien des Pjnrendes. Bull. Soc. geol. Fr., 

Paris (7) 3 : 437-448, text-plate. 
Trauth, F. 1918. Das Eozanvorkommen bei Radstadt im Pongau. Denkschr. Akad. Wiss. 

Wien, 95 : 171-278. 
Weber van Bosse & Foslie, M. H. 1904. Corallinaceae Siboga Expedition. Siboga Exped., 

Leiden, 61 : i-iio, pis. 1-16. 



PLATE I 

(All X50) 

Fig. I. Archaeolithothamnium lauensum Johnson & Ferris. Vertical section of a 
fertile fragment. Lower Miocene. V. 51758. 

Fig. 2. Archaeolithothamnium lugeoni Pfender. A somewhat oblique section. Paleo- 
cene. V.51760. 

Figs. 3, 4. Archaeolithothamnium aschersoni (Schwager). 3, Vertical section of a thin 
crust. V.51762. 4, Oblique section of a small rounded knob. Paleocene. V.51761. 

Fig. 5. Archaeolithothamnium cf. cyrenaicum Raineri. A slightly oblique section. 
Middle Oligocene. V. 51 759. 

Fig. 6. Archaeolithothamnium intermedium Raineri. A nearly vertical section. 
I^ower Miocene. V. 51758. 



Bull.BM. {N.H.) Geol. 11,6 



PLATE 1 




PLATE 2 
(All X50) 

Fig. I. Archaeolithothamnium saipanense Johnson. Vertical section. Middle Oligocene. 

V.5I763- 

Fig. 2. Lithothamnium cantabricum Lemoine. Vertical section showing hypothallus 
and perithallus. Paleocene. V.51766. 

Fig. 3. Archaeolithothamnium sociabile Lemoine. Nearly vertical section. Middle 
Oligocene. V. 51 764. 

Fig. 4. Lithothamnium cf . caucasicum Maslov. Slightly oblique section of a small branch. 
Paleocene. V.51767. 

Fig. 5. Archaeolithothamnium sarawakense n. sp. Holotype. Vertical section. Paleocene. 
V.51762. 

Fig. 6. Lithoporella melobesioides (Foslie). Paleocene. V.51766. 



Bull.B.M. (N.H.) Geol. 11, 6 



PLATE 2 




PLATE 3 
(All X50) 

Figs, i, 2. Mesophyllutn cf. pfenderae (Lemoine). Sections of small branches. Fig. 2 
shows a conceptacle with sporangia. Paleocene. V. 51768. 

Fig. 3. Mesophyllutn curtum Lemoine. Vertical section of a branch, badly recrystallized. 
Lower Miocene. V. 51769. 

Fig. 4. Archaeolithothamnium macrosporangium n. sp. Holotype. Vertical section 
showing tissue and sporangia. Lower Oligocene. V. 51 765. 

Fig. 5. Lithoporella cf. minus Johnson. Badly recrystallized. A conceptacle chamber 
to right. Paleocene. V.51760. 

Figs. 6, 7. Corallina cf. abundans Lemoine. Fig. 6 shows a terminal conceptacle 
chamber. V. 51778. Fig. 7, Fragment of a segment. Lower Oligocene. V.51779. 



BuU.B.M. {N.H.) Geol. 11, 6 



PLATE 3 






PLATE 4 
(All X50) 

Figs, i, 6. Mesophyllum vaughanii (Howe) var. sarawakense nov. Fig. i, A nearly 
vertical section, Holoty|De. Middle Oligocene. V. 51772. Fig. 6, An oblique section. Middle 
Oligocene. V. 51759. 

Fig. 2. Jania vetus Johnson. Lower Miocene. V. 51758. 

Fig. 3. Mesophyllum vaughanii (Howe). Lower Oligocene. V.51771. 

Fig. 4. Lithophyllum quadrangulum Lemoine. Section of a long thin crust mainly 
hypothallus. Middle Oligocene. V.5177^. 

Fig. 5. Lithophyllum cf. obliquum Lemoine. Vertical section. Miocene. V. 51773. 



Bull.B.M. {N.H.) Geol. 11, 6 



PLATE 4 




li 


^^M 






2 


W^' 


^^^ 


l'^x'^^ 




fl 


m 




1 


-i 1 f^^^^ 


1 


^ 
V 


"m^ 


r rt VVjS 




H 


4 


«l 




^^.>k'-v.> 


■s ^ . 



4 





1 



PLATE 5 

Fig. I. " Solenomeris" sp. Vertical section of a fragment, X40. Paleocene. V. 51784. 

Fig. 2. Pycnoporidium sinuosum Johnson & Konishi (X50). Paleocene. V. 51785. 

Fig. 3. Jania miocenica Johnson (X50). Lower Miocene. V.51758. 

Fig. 4. Corallina neuschelorum Johnson. A partial segment, X50. Lower Miocene. 
V.51758. 

Fig. 5. Mesophyllum vaughanii (Howe). Fragment showing conceptacle chambers, 

X50. Lower Oligocene. V. 51770. 

Fig. 6. Lithophyllum capederi Lemoine. Several crusts, X50. Lower Miocene. 

V.5I775- 

Fig. 7. Distichoplax biserialis (Dietrich). Sections of two fragments, X40. Paleocene, 
Borneo. V.51783. 

Fig. 8. Lithophyllum densum l^emoine. Slightly oblique long section of a branch, X50. 
Paleocene. V. 51767. 



BiM.B.M. {N.H.) Geol. 11,6 



PLATE 5 




PLATE 6 

(All X50) 

Figs, i, 2. Subterraniphyllum thomasi Elliott. Fig. i, An oblique long section. Lower 
Oligocene. V. 51782. Fig. 2, Longitudinal section of a worn fragment. Lower Oligocene. 
V.51782. 

Fig. 3. Lithoporella antiquitas Johnson. Two conceptacle chambers shown. Lower 
Miocene. V. 51776. 

Fig. 4. Dermatolit Hon saipanense ]ohnson. Lower Miocene. V.51758. 

Fig. 5. Jania cf. nummulitica Lemoine. Lower Oligocene. V. 51780. 

Fig. 6. Thaumatoporella parvovesiculif era (Ruinen). Paleocene. V. 51777. 

Fig. 7. Amphiroa cf. fortis Johnson. Slightly oblique vertical section. Paleocene. 
V.51781. 



Bull. B.M. (N.H.) Geol. 11,6 



PLATE 6 





^Sii v^\C 



PRINTED IN GREAT BRITAIN 
BY ADLARD & SON LIMITED 
BARTHOLOMEW PRESS, DORKING 



BRITISH WEALDEN SHARKS 



C. PATTERSON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. ii No. 7 

LONDON: 1966 



BRITISH WE ALDEN SHARKS | i ^ jan 19 



BY 



COLIN PATTERSON, Ph.D. 



Pp. 281-350; 5 Plates; 31 Texi-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

GEOLOGY Vol. 11 No. 7 

LONDON: 1966 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 7 of the Geological 
[Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 



Trustees of the British Museum (Natural History) 1966 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued II January 1966 Price £2 2s. 



BRITISH WEALDEN SHARKS 

By COLIN PATTERSON 



CONTENTS 

I Introduction 
II Localities 

III Systematic descriptions 

Class Selachii 

Order Hybodontiformes 
Family Hybodontidae 

Hybodus basanus Egerton 

Hybodus ensis Smith Woodward 

Hybodus parvidens Smith Woodward 

Hybodus brevicostatus sp. nov. . 

Fin spines of Hybodus 

Remarks on hybodont teeth 

Lonchidion breve breve sp. et ssp. nov. 

Lonchidion breve crenulatum ssp. nov. 

Lonchidion breve pustulatum ssp. nov. 

Lonchidion striatum sp. nov. 

Lonchidion rhizion sp. nov. 

Lonchidion heterodon sp. nov. 

Fin spines and cephalic spines of Lonchidion 

The Affinities of Lonchidion 
Family Ptychodontidae 

Hylaeobatis ornata (Smith Woodward) 

The Afi&nities of Hylaeobatis 

The origin of Hylaeobatis 

Relationships within the Ptychodontidae 

IV ecology and relationships of the fauna 
V References ...... 



Page 
284 

285 
287 
287 
287 
287 
288 
292 
296 
300 
310 
311 
313 
316 
317 
320 
322 
326 
328 
330 
332 
333 
339 
340 
342 
346 
348 



SYNOPSIS 

New material, consisting mainly of abundant isolated teeth from bone-beds, allows more 
detailed and precise definition of the known sharks of the British Wealden and Purbeck, and 
contains five new species : Hybodus brevicostatus , Lonchidion breve (with three new subspecies), 
L. striatum, L. rhizion and L. heterodon. Samples from successive horizons allow the inter- 
relationships of the various species to be worked out in some detail. Hylaeobatis is shown to 
belong in the family Ptychodontidae and to lie near the ancestry of Ptychodus : the Ptycho- 
dontidae are probably derived from the hybodontid genus Lonchidion. The Wealden shark 
fauna is unusual in being from fresh water and in containing only hybodontoids : it is suggested 
that the hybodonts were able to escape from competition with more advanced selachians by 
entering fresh water : the radiation which they underwent there parallels their marine radiation 
at their first appearance, and explains the similarity between the shark fauna of the Wealden 
and of the marine Triassic. Certain of the more specialized Wealden forms seem to have returned 
to the sea and given rise to the Upper Cretaceous hybodonts and ptychodonts. 



GEOL. II, 7 



27 



284 BRITISH WEALDEN SHARKS 

I INTRODUCTION 

This paper has been prompted by the arrival in the British Museum (Natural History) 
of two sets of new Wealden material. Between i960 and 1962 Drs. W. A. Clemens 
and K. A. Kermack of University College, London, were searching the bone-beds 
of the British Wealden for mammalian remains. In the course of this work large 
quantities of bone-bed were broken down with formic acid, and the bone separated 
by bromoform flotation. The mammalian finds resulting from the work have 
already been described (Clemens i960, 1963; Kermack, Lees & Mussett 1965), and 
Dr. Kermack and his colleagues were kind enough to present the residue of the 
treated bone-beds to this museum. This material is rich in fish remains, though the 
majority are rolled and water-worn. In ig6i Mr. J. F. Wyley of Richmond, Surrey, 
discovered a bone-bed in the Weald Clay in the Henfield Brick Company's pit at 
Henfield, Sussex, and in many visits to the pit he has collected a quantity of fish 
material which he has generously presented to this museum. The material from 
Henfield is normally excellently preserved. 

Knowledge of British Wealden sharks is due almost entirely to Smith Woodward. 
Agassiz (1837) described several hybodont fin spines under various names, and 
Egerton (1845) described Hyhodus hasanus from the Weald Clay of the Isle of Wight 
and (1854) 3- fi^ spine from Tilgate as Asteracanthus granulosus. In 1889 Smith 
Woodward briefly redescribed these species, referred several teeth to Hyhodus, 
and described Acrodus ornatus from the Upper Wealden. Later (Smith Woodward 
1916), in his monograph on the fishes of the British Wealden and Purbeck, he gave 
more detailed descriptions of all these species and added two new species of Hyhodus, 
H. ensis and H. parvidens, and a new genus and species, Hylaeohatis prohlematica, 
all based on isolated teeth. No new material of any of these species has since been 
described [H. hasanus has been incorrectly recorded from the Cretaceous of Japan 
by Yabe & Obata 1930). 

In this paper a new species of Hyhodus and four new species, one with three sub- 
species, of the hybodont genus Lonchidion Estes (1964) are described, and Acrodus 
ornatus and Hylaeohatis prohlematica are shown to be synonymous. The total list 
of British Wealden sharks known at present is therefore : 

Hyhodus hasanus Egerton 
*H. ensis Smith Woodward 
*H. parvidens Smith Woodward 

H. hrevicostatus sp. nov. 
f/f . striatulus Agassiz 
^Lonchidion hreve sp. nov. 
*L. striatum sp. nov. 
*L. rhizion sp. nov. 
*L. heterodon sp. nov. 

-f Asteracanthus granulosus Egerton (doubtfully Asteracanthus, see p. 310) 
*Hylaeohatis ornata (Smith Woodward) 

* Species known only by isolated teeth, 
t Species known only by isolated fin spines. 



BRITISH WEALDEN SHARKS 285 

II LOCALITIES 

The bulk of the new material described here is from four horizons, the Cliff End 
bone-bed, the Telham bone-bed, the Paddockhurst bone-bed and the Weald Clay of 
Henfield, Sussex. 

{a) Cliff End Bone-bed 

The Cliff End bone-bed, exposed on the foreshore at Cliff End, near Hastings, 
Sussex, the source of the mammal teeth discovered early in the century by Teilhard 
de Chardin and Pelletier, has recently been described by Allen (i960 : 11) and Clemens 
(1963 : 58), the latter also describing the method used to concentrate the vertebrate 
remains. The bone-bed is within the Ash down Beds (of Valanginian age according 
to Hughes 1958), but its precise horizon is not yet established*. The Ashdown Beds 
are interpreted by Allen (1959) as a deltaic deposit, the delta flowing into a fresh- 
water lake. The great majority of the vertebrate remains are strongly rolled and 
abraded. Of the recognizable fragments about 45% are shark teeth, 50% are teeth 
and fragments of dermal bones of actinopterygians (mainly Lepidotes), and 5% or 
less are reptilian. 

The shark fauna includes : 

Hybodus ensis (common) 

H. parvidens (common) 

H. brevicostatus (rare) 

Lonchidion breve breve (moderately common) 

L. rhizion (uncommon) 

L. heterodon (rare) 

(b) Telham Bone-bed 

The Telham bone-bed is exposed at a number of localities in the south-eastern 
Weald (Allen 1949 : 279, text-fig. 45). The sample described here was collected by 
Mr. P. J. Whybrow from the exposure at Teigh Farm, Stone, Kent (GR TQ 937268). 
The bone-bed lies in the Wadhurst Clay, near the base (Upper Valanginian according 
to Hughes 1958), and is interpreted by Allen as the result of river water flooding over 
a delta plain. The vertebrate remains are in much the same condition as at Cliff 
End, rolled and abraded. Of the recognizable fragments, about 30% are shark 
teeth and the remaining 70% are almost entirely actinopterygian, mainly Lepidotes : 
reptiles account for only about 1% of the sample. The shark fauna includes : 

Hybodus ensis (rare) 

H. parvidens (common) 

Lonchidion breve breve (moderately common) 

L. rhizion (rare) 
This locality is referred to in the text as Telham bone-bed. Stone, Kent. 

* In a paper published while this work was in press, Kermack, Lees & Mussett (1965 : 536) give further 
information on the Cliff End bone-bed, pointing out that the bed cannot now be located in the cliff or 
on the foreshore, and that the scattered blocks found on the beach probably come from an off-shore 
reef, which would place the bone-bed "well down in the Fairlight Clays", near the base of the Wealden. 
Kermack, Lees & Mussett also publish a comment on the Cliff End fauna which I wrote on first seeing 
the material. I would not now infer brackish conditions from the abundant hybodont selachians : 
I think it probable that all the species present were capable of life in fresh water. 



286 BRITISH WEALDEN SHARKS 

(c) Paddockhurst Bone-bed 

Clemens (i960, 1963 : 63) has given this name to a bone-bed in the Grinstead Clay 
(Hauterivian according to Hughes 1958) at Paddockhurst Park, near Turner's Hill, 
Sussex. Allen (1959) interprets the Grinstead Clay as a fresh-water lake deposit. 
The vertebrate remains in the Paddockhurst bone-bed are neither so broken up nor 
so badly rolled as they are at Cliff End, and sharks are less common : shark teeth 
make up about 15% of the recognizable elements, reptiles about 10%, and the 
remaining 75% is actinopterygian, mainly scales and teeth of Lepidotes. 
The shark fauna includes : 

Hybodus ensis (common) 

H. parvidens (common) 

H. brevicostatus (rare) 

Lonchidion breve breve (moderately common) 

L. breve crenidatum (moderately common) 

L. rhizion (rare) 

{d) Henfield 

The Henfield Brick Company's pit (GR TQ/218143) at Henfield, Sussex, is a large 
pit in the Weald Clay which is being actively worked (Milbourne 1961 : 135). The 
beds dip northwards at about 5 degrees (Reeves 1947 : 83). The highest beds 
exposed, in the north wall of the pit, are red and yellow clays without obvious fossils. 
These beds pass down into brown and grey clays. About 3 ft. below the red and 
yellow clays there is a band of ' Paludina ' limestone, 3-8 in. thick. Fishes occur in 
the clay immediately above the limestone. Below the limestone there is 10 ft. of 
brown sandy clay, and below this about 25 ft. of grey clay in which six minor cyclo- 
thems, 3-4 ft. thick, are recognizable by increasing sand content giving a brownish 
tinge towards the top of each. Each of these cyclothems contains fish fragments, 
often concentrated into more or less well marked bone-beds which are occasionally 
cemented into a phosphatic limestone (PI. i, fig. i). Below these cyclothems the 
floor of the pit is occupied by grey clay containing Iguanodon, lignite and occasional 
tree trunks, passing down in the south wall of the pit into barren red and yellow 
clays and sands, the lowest beds exposed. This sequence lies in Reeve's Group H 
(1958 : 11), the middle division of the Weald Clay (Barremian according to Allen 
1955 and Hughes 1958), probably near the top : Dr. F. W. Anderson has examined 
samples of ostracods collected from the various fish horizons and places the ' Paludina ' 
limestone at about 550 ft. below the top of the Weald Clay. 

Associated with the fishes there are abundant ostracods, all non-marine according 
to Dr. Anderson, and occasional charophyte oogonia, another indication of fresh water 
conditions. The fish material, although consisting almost entirely of dissociated 
fragments, is well preserved. There is no significant difference in the fish fauna 
of the various fish horizons, but in the minor cyclothems the fauna varies widely 
within the bone-beds: the block of bone-bed shown in PI. i, fig. i shows abundant 
shark teeth of five species (teeth of Lonchidion breve and L. striatum are too small 
to show on the photograph), but the other side of the block, which is only 14 mm. 



BRITISH WEALDEN SHARKS 287 

thick, shows only two or three teeth of Lonchidion, and is made up ahnost entirely of 
teleostean remains. Sharks make up less than io% of the material, the bulk of 
which is teleostean. Actinopterygians identified include : 

Lepidotes mantelU Agassiz 

Coelodus mantelli Agassiz 

Caturus tenuidens Smith Woodward (PI. i, fig. 2 : not previously recorded 
above the Purbeck) 

Pachythrissops sp. 

Clupavus sp. 
The shark fauna includes : 

Hybodus basanus (common) 

H. parvidens (rare) 

H. brevicostatus (moderately common) 

Lonchidion breve breve (common) 

Lonchidion striatum (common) 

Hylaeobatis ornata (common) 
[e) Other localities 

Small but valuable samples from two other locaUties are also described here. 
The first was collected by Mr. I. M. West of the University of Southampton from a 
limestone above the Broken Shell Limestone in the Upper Purbeck exposed loo yards 
east of Friar Waddon farm, near Upwey, Dorset (GR SY/643858) (see Anderson 
1958 : iig, 129, text-figs. 21, 22). Mr. West thinks this horizon is equivalent to the 
Unio Bed of the type Purbeck. The second sample was collected by Mr. P. J. 
Whybrow from a limestone in the Wadhurst Clay exposed in a cutting on the east 
side of the road 200 yards north of Homeland, Ashurstwood, Sussex (GR TQ/419363). 
These two localities are referred to in the text as Friar Waddon and Ashurstwood 
respectively. 

Ill SYSTEMATIC DESCRIPTIONS 

Class SELACHII 
Order HYBODONTIFORMES 
Family HYBODONTIDAE Owen 1846 
Diagnosis. See Berg (1955 : 61) 

Genus HYBODUS Agassiz 1837 : 41 
Diagnosis. See Smith Woodward (1916 : 3), but delete ' palatoquadrate not 
articulated with the postorbital region of the skull '. 
Type species. Hybodus reticulatus Agassiz. 

H. basanus Egerton, H. ensis Smith Woodward and H. parvidens Smith Woodward, 
the three species of the genus known by teeth in the British Wealden, are easily 
distinguished in the type material described by Smith Woodward but appear to be 
linked by intermediate forms in the new material. H. basanus is the only species of 
the three in which the complete dentition is known, and will be described first. 



288 BRITISH WEALDEN SHARKS 

Hybodus basanus Egerton 
(PI. I, fig. I ; Text-figs. 1-3) 

1845 Hyhodiis basanus Egerton : 197, pi. 4. 

1889 Hybodus basanus Egerton ; Smith Woodward : 273, pi. 12, figs. 1-5. 

1891 Hybodus basanus Egerton ; Smith Woodward : 63, pi. i, pi. 2, fig. i. 

1898 Orthybodus basanus (Egerton) Jaekel : 139. 

1916 Hybodus basanus Egerton ; Smith Woodward : 5, pi. i, figs, i, 2 ; pi. 2, fig. i ; text-figs. 

3-5- 
1919 Hybodus basanus Egerton ; Smith Woodward : 139, pi. 26, fig. 3. 

Amended diagnosis. Hyhodus known by skull, dentition, fin spines and frag- 
ments of postcranial skeleton ; nine or ten files of teeth in each ramus of the jaws, 
symphysial file present in lower jaw, dentition weakly heterodont ; teeth reaching 
15 mm. in length, central cusp high, slender (ratio of height above root/crown 
junction to length i-o-2-o) and arched lingually ; three or (rarely) four pairs of 
lateral cusps ; labial face of crown with rather numerous fine striae, often bifurcated 
basally, reaching tips of lateral cusps and covering from one-third to two-thirds of 
central cusp ; lingual face of crown with similar but longer striae almost reaching 
tip of central cusp ; teeth without accessory cusps on the labial or Ungual margins ; 
root low, bent linguaUy ; a single pair of cephalic spines with terminal barb ; fin 
spines slender, compressed, not much arched, reaching about 20 cm. in length, 
lateral faces with 8-12 fine, sharp ridges, posterior denticles in two series, small and 
closely set. 

HoLOTYPE. GSM No. 27973, Geological Survey & Museum, London, from the 
Weald Clay of Atherfield, Isle of Wight. 

Material. In addition to the holotype, about twenty skulls, fragments of 
vertebral column, fin spines and isolated teeth in the British Museum (Natural 
History) and the Geological Survey & Museum. 

Horizon and localities. Weald Clay : Atherfield, Isle of Wight ; Pevensey 
Bay, Sussex ; Henfield, Sussex ; Bexhill, Sussex. 

Description. Hybodus basanus is the only species of the genus in which the 
skull is well known. Smith Woodward (1916) has given a good description of the 
skull based on several well preserved heads in the British Museum (Natural History) . 
Of the palato-quadrate he writes ' it can scarcely have articulated with the post- 
orbital prominence of the neurocranium '. Smith (1942 : 701) has questioned this, 
noting that in the Liassic H. hauffianus Fraas the skull is said to be amphistylic by 
Jaekel, and that it is amphistylic in Synechodus. Berg (1955 : 62) has also suggested 
that Smith Woodward's observation needs checking. I have examined all of the 
skulls of H. basanus in the British Museum (Natural History) . The palatoquadrate 
and hyomandibular are best shown in P. 2082a and P. 6103. There can be no doubt 
that Smith Woodward's reconstruction (1916, text-fig. 3) is accurate in showing the 
hyomandibular as being large and much higher than the postorbital part of the 
palatoquadrate. In this H. basanus differs from typical amphistylic sharks such as 
Hexanchus, where the hyomandibular is slender and no higher than the otic process 



BRITISH WEALDEN SHARKS 



289 






2 mm 



Fig. I. Hybodus basanus Egerton. Tooth from the first or second file of the upper jaw, right 
side, in labial (a), lingual (b) and medial (c) view. P.11871, Weald Clay; Pevensey Bay, 
Sussex. 



of the palatoquadrate, and from Heterodontus and Hybodus hauffiamis (Jaekel 1906) 
in which the hyomandibular is stout but equal in height to the otic process of the 
palatoquadrate. I do not think it is possible, however, to decide whether the 
palatoquadrate of H. basanus had a post-orbital articulation or not. The otic 
process was certainly tucked well up below the postorbital process of the neuro- 
cranium, but there is no sign of an articular facet or condyle on the otic process. 



290 



BRITISH WEALDEN SHARKS 



It is possible that the suspension was amphistyhc, but the long, large hyomandibular 
shows that the species tends towards the hyostylic condition. 

Smith Woodward describes the dentition of H. basanus as consisting of ten or 
eleven paired files of teeth in each jaw, with a symphysial file in the mandible. 
The teeth (Text-fig. i) have a high, slender central cusp and three pairs of lateral 
cusps which are well separated from the central cusp. The height of the central 
cusp (above the root /crown junction) is only slightly less than the length of the tooth 
in anterior teeth, but the height decreases posteriorly. 

The central cusp curves lingually rather strongly, but the tip curves labially again, 
giving a weakly sigmoid outline to the cusp in medial view (Text-fig. ib). Of the 
three pairs of lateral cusps, the innermost is sharply pointed, striated to its tip, and 
normally about one-third as high as the central cusp. The second pair of lateral 
cusps is shorter but similarly ornamented. The outermost cusps are normally very 
small and almost smooth. A minute fourth lateral cusp is occasionally present. 





2 mm 




Fig. 2. Hybodus basanus Egerton. a. Tooth from the first file of the upper jaw, right side, 
in labial view. P. 2082. b. Tooth from the sixth file of the lower jaw, left side, in labial 
view. P.6356. c. Tooth from the eighth file of the lower jaw, left side, in labial view. 
P.20826. All from the Weald Clay of Pevensey Bay, Sussex. 



BRITISH WEALDEN SHARKS 



291 



The base of the labial face of the crown is ornamented with moderately fine, 
parallel or sub-parallel vertical striae, often bifurcating basally, which extend to the 
tips of the lateral cusps and to about one-third or half of the height of the central 
cusp. There is a good deal of variation in the strength, number and length of these 
striae : they are closely packed, weak and short in some fish (P.2082&, Text-fig. 
2C), and coarse, well spaced and long, covering more than half the labial face of 
the central cusp, in others (P.11871, Text-fig. ia). On the lingual face of the 
crown the ridges are stronger, and extend almost to the tip of the central cusp 
(P. 2082a, P.11871, Text-fig. IB). The root is low, and is turned lingually, with 
the labial face of the root lying almost at right angles to the axis of the central cusp. 
This description is based on the teeth of the complete skulls from the Weald Clay of 
Pevensey Bay, Sussex, and Atherfield, Isle of Wight. 

It is difficult to match these teeth exactly in material from any other Wealden 
horizon or locality. In the new material from the Weald Clay of Henfield, which 
is approximately contemporary with the type material of H. hasanus, there is a 
number of teeth of the type shown in PL i, fig. i and Text-fig. 3. In these teeth 
the crown is lower than in those from the complete skulls of H. basanus, with a ratio 
of height (above the root/crown junction) to length of little less than 2-0, the striae 
are less numerous and are longer, reaching well over half the height of the central 




2 mm 



Fig. 3. Hybodiis basanus Egerton. Lateral tooth in lingual (a), distal (b) and labial (c) view. 
P. 4692 1, Weald Clay ; Henfield, Sussex. 



292 BRITISH WEALDEN SHARKS 

cusp on the labial face (the tooth shown in Text-fig. 3 has shorter striae than most 
teeth from Henfield) and to the tip of the cusp on the lingual face, and both the 
central cusp and the root are not turned lingually so strongly as they are in typical 
H. basanus (cf. Text-figs, ic, 3c). In all these features the teeth from Henfield are 
intermediate between typical H. basanus from Atherfield and Pevensey, and typical 
teeth of H. parvidens from the Ashdown Beds and Wadhurst Clay (Text-figs. 6, 7). 
The Henfield teeth are referred to H. basanus rather than H. parvidens because they 
never show the knob or accessory cusp at the base of the labial face of the central 
cusp which is common in H. parvidens, because the striae are almost as numerous as 
they are in H. basanus and only rarely reach the tip of the central cusp, and because 
there is a difference in the extent of the striae on the labial and lingual faces of the 
central cusp (in H. parvidens the length of the striae is similar on the two faces of 
the central cusp) . 

No teeth referable to H. basanus are known from horizons above or below the 
Weald Clay. The tooth from the Lower Cretaceous of Japan referred to H. basanus 
by Yabe & Obata (1930 : 4, pi. 2, fig. 3) differs from this species in the longer and more 
slender central cusp and in the almost complete lack of striae on the labial face : 
it is possibly a Synechodus. Leriche (1911 : 457, pi. 6, fig. 2) has referred to H. 
basanus a fragment of fin spine from the Lower Neocomian of the Paris Basin, but 
this identification is very doubtful : the fragment could as well belong to another 
species. 

Affinities. Smith Woodward (1916 : 10) considered that teeth and spines from 
horizons in the lower part of the Wealden showed resemblances to H. basanus without 
being referable to the species. These forms (see p. 294) are here included in H. ensis 
which in the Wealden seems to trend towards H. basanus in the form of the teeth, 
but is probably not related. Teeth of H. basanus from Henfield are intermediate 
in a number of characters between more typical teeth of the species from Pevensey 
and the Isle of Wight, and teeth of H. parvidens from lower horizons. This, together 
with the trend towards H. basanus shown by H. parvidens in the Grinstead Clay 
(see p. 299), leaves little doubt that H. basanus is a species confined to the Weald 
Clay (probably to the upper part alone) which evolved directly from H. parvidens 
by increase in size, in crown height and in the number of striae. 

Hybodus ensis Smith Woodward 
(Text-figs. 4, 5) 

1889 Hybodus sp. inc. (? styictus Agassiz) Smith Woodward : 275. 

1889 Hybodus sp. inc. (? striatulus Agassiz) Smith Woodward : 276, pi. 11, figs. 14, 15. 

1916 Hybodus ensis Smith Woodward : 11, pi. 2, figs. 2-6; non fig. 7. 

Amended diagnosis. Hybodus known only by isolated teeth : teeth large, up to 
2 cm. in length, central cusp high (ratio of height above root /crown junction to 
length I-0-I-5), broad at base, evenly tapering and moderately compressed, not much 
arched lingually ; two pairs of lateral cusps, inner lateral cusps moderately high, 
slender, pointed, well marked off from central cusp but close to it ; labial face of 



BRITISH WEALDEN SHARKS 



293 



crown with many fine, parallel striae reaching tips of lateral cusps but covering only 
basal quarter or fifth of central cusp ; lingual face of crown with coarser striae 
extending about half way up central cusp ; root low, bent lingually. 

HoLOTYPE. BMNH No. 21349 (Smith Woodward, 1916, pi. 2, fig. 6), tooth 
without root. Middle Purbeck ; Swanage, Dorset. 

Material. In addition to the holotype, about one hundred isolated and frag- 
mentary teeth. 

Horizons and localities. Middle Purbeck : Swanage, Dorset. Upper Purbeck : 
Friar Waddon, Dorset. Ash down Beds : Cliff End, Sussex. Wadhurst Clay : Teigh 
Farm, Stone, Kent ; Hastings, Sussex. Grinstead Clay : Paddockhurst Park, 
Sussex ; Tilgate Forest, Sussex. 

Description. Smith Woodward based this species on incomplete teeth from the 
Middle Purbeck of Dorset, but also recorded it from the Wealden of Tilgate Forest 
(Grinstead Clay — see p. 294). All the specimens labelled as H. ensis by Smith 
Woodward are from the Purbeck. 




Fig. 



Hybodus ensis Smith Woodward. Tooth in labial (a), lingual (b) and medial (c) 
view. 21349c, Middle Purbeck ; Swanage, Dorset. 



294 BRITISH WEALDEN SHARKS 

The species is poorly known, the Purbeckian type material consisting of teeth 
without roots exposed in labial view. One of these teeth, 21349c, has been removed 
from the matrix to expose the lingual face (Text-fig. 4). In the Purbeck material 
of H. ensis the crown is about as high as in H. basanus, with the ratio of length to 
height (above the root/crown junction) about i-o, but the central cusp is much 
broader, the ratio of the breadth at its middle point to its height ranging from 
2-8 to 3-0, compared with 4-0 to 5-0 in H. basanus. In no specimen of H. ensis 
are there more than two pairs of lateral cusps (21349&, Smith Woodward 1916, pi. 2, 
fig. 7, has three pairs of lateral cusps but does not belong in H. ensis — see H. parvidens, 
p. 297). In H. ensis the labial face of the crown (Text-fig. 4A) bears fine, close 
packed, parallel striae which extend almost to the tips of the inner lateral cusps but 
cover only the basal one-fifth or quarter of the central cusp. On the lingual face of 
the crown (Text-fig. 4B) the striae are less numerous, a little coarser and cover almost 
one-third of the height of the central cusp. The central cusp curves lingually less 
strongly than it does in H. basanus (cf. Text-figs, ic, 4c). Other apparent differ- 
ences between H. ensis and H. basanus suggested in Smith Woodward's original 
description — that the inner lateral cusps are higher and closer to the central cusp in 
H. ensis, and that the ratio of height to length of the tooth is greater in this species — 
are not confirmed by measurement of the teeth. 

In 1889 (p. 276) Smith Woodward catalogued as ' Hybodus sp. inc. (? striatulus 
Agassiz) ' about thirty teeth from the Wealden of Tilgate Forest,^ noting the high, 
moderately broad central cusp, the high, slender inner lateral cusps, and the striae 
' rarely extending more than half the height of the median cone '. In 1916 (p. 10) 
he referred to these teeth as ' of the same general type as those of H. basanus . . . but 
not sufficiently similar to be referred with certainty to this species '. Most of these 
teeth are more or less rolled and waterworn, and many are quite indeterminable. 
Some, such as 48377 (Tilgate Forest) and P. 6353 (Hastings, unknown horizon) 
agree exactly with the Purbeck specimens of H. ensis, and leave no doubt that this 
species extended into the Wealden. But the majority of these teeth do not agree 
exactly with the type material of H. ensis : 2693 (Tilgate Forest, Text-fig. 5, Smith 
Woodward 1889, pi. 11, fig. 14) is the best preserved of these, and is typical in shape. 
These teeth differ from H. ensis in the more slender, sharply pointed central cusp 
(ratio of breadth measured at the middle point to height c. 3-5), but they agree with 
H. ensis and differ from H. basanus in their large size, weakly arched central cusp 
(Text-fig. 5b), two pairs of lateral cusps, in the short, fine striae on the labial face of 
the crown, rarely reaching more than one-third of the height of the central cusp, 
and in the short striae on the lingual face of the crown. The striae on the lingual face 
of the central cusp extend about as far as they do in typical H. ensis, and are charac- 
teristically longer near the margins of the cusp than they are in the centre. In my 
opinion these teeth should be included in H. ensis. Some of these Wealden teeth 
(26026 ; Smith Woodward 1889, pi. 11, fig. 14 : 26024, both from Tilgate Forest) 

1 Mantell's horizon at Tilgate Forest, previously thought to lie in the Upper Tunbridge Wells Sand, 
directly below the Weald Clay (Topley 1875 : 92), has recently been re-identified as in the Cuckfield 
Stone, in the middle part of the Grinstead Clay (Stubblefield 1963 : 37). 



BRITISH WEALDEN SHARKS 



295 




2 mm 

Fig. 5. Hyhodus ensis Smith Woodward. Tooth in labial (a) and medial (b) view. 
2693, Grinstead Clay ; Tilgate Forest, Sussex. 



tend towards H. basanus in having a more slender central cusp with rather coarse 
striae covering almost half of the labial face, but they have only two pairs of lateral 
cusps : 26024 is particularly like the typical Weald Clay H. basanus, but it seems 
probable that it is not an early example of this species but a tooth of H. ensis in 
which the evident trend towards narrowing of the central cusp and extension and 
coarsening of the ornament has gone farther than in other examples. 

Among the new material from the Cliff End and Paddockhurst bone-beds there 
are no complete large teeth, but isolated central cusps and imperfect crowns are 
very common. All of these large teeth can be referred to H. ensis : they show 
a range in breadth of the central cusp from stout forms like the Purbeck H. ensis 
to slender forms approaching the proportions of H. basanus, but the striae on the 
labial face are always short, fine and closely packed, and on the lingual face they are 
always shorter than in H. basanus and are longer at the margins of the central cusp 
than in the centre. 

No examples of H. ensis are known from horizons above the Grinstead Clay. 

Affinities. H. ensis is a species which ranges from the Middle Purbeck to the 
Grinstead Clay (Paddockhurst bone-bed and Tilgate Forest) . In the Wealden the 
teeth show a trend towards narrowing of the central cusp, sometimes accompanied 
by coarsening and lengthening of the striae which may produce teeth closely similar 
to those of H. basanus in shape. 



296 BRITISH WEALDEN SHARKS 

Smith Woodward (1916 : 11) noted the similarity between the Purbeck forms of 
H. ensis and the Middle and Upper Jurassic H. grossiconus Agassiz. Extremely 
similar to H. ensis is H. songaensis Saint-Seine (1962 : 4, pi. 6, fig. 6), a species based 
on a single tooth from the Songa Beds of the Congo, probably marine and of Kim- 
meridgian age, which could well be synonymous with H. ensis. H. ensis is possibly 
derived directly from these marine Jurassic forms. 

Hybodus parvidens Smith Woodward 
(Text-figs. 6-9) 

1889 Hybodus sp. inc. Smith Woodward : 276, pi. 11, fig. 16. 

1916 Hybodus ensis Smith Woodward : pi. 2, fig. 7 (errore). 

1916 Hybodus parvidens Smith Woodward : 12, pi. 2, figs. 8-14. 

1949 Hybodus parvidens Smith Woodward ; Allen : 277 (name only). 

Amended diagnosis. Hybodus known only by isolated teeth : teeth small, less 
than 10 mm. in length ; central cusp moderately high, not much copipressed, lower 
and broader in posterior teeth ; ratio of height of central cusp (above root /crown 
junction) to length of tooth between 1-5 and 3-0 ; three or four pairs of lateral cusps ; 
labial face of crown with few coarse striae, often bifurcating basally, which reach 
the tips of the lateral cusps and commonly reach the tip of the central cusp except 
in some high anterior teeth, striae on lingual face similar ; knob or accessory cusp 
frequently present at base of labial surface of central cusp, no other accessory cusps ; 
root moderately deep, turned lingually a little. 

HoLOTYPE. BMNH P.11877, tooth without root, Wadhurst Clay, Hastings. 

Material. About two hundred isolated teeth. 

Horizons and localities. Middle Purbeck : Swanage, Dorset. Upper Purbeck : 
Friar Waddon, Dorset. Ashdown Beds : Cliff End, Sussex ; Fairlight, Sussex. 
Wadhurst Clay : Teigh Farm, Stone, Kent ; Hastings, Sussex ; Rye, Sussex. 
Grinstead Clay : Paddockhurst Park, Sussex ; Tilgate Forest, Sussex. Weald Clay : 
Henfield, Sussex. 

Description. Smith Woodward based this species on incomplete teeth from the 
Wadhurst Clay, in none of which was the root preserved. The holotype and two 
of the paratypes are shown in Text-fig. 6. Among the new material from the Cliff 
End bone-bed there are several more or less complete and unworn teeth (Text-fig. 7) . 
The high-crowned teeth (Text -figs. 6, 7A) are presumably anterior, the low-crowned 
(Text-fig. 7b) posterior. 

In this typical material from the Ashdown Beds and Wadhurst Clay, the central 
cusp is moderately high in anterior teeth, with a ratio of height of cusp to length of 
tooth of about I-8-2-0. In posterior teeth the central cusp becomes much lower, the 
ratio of height to length reaching about 3-0. The central cusp is not compressed as it 
is in H. ensis and H. basanus, and does not curve lingually. There are always three 
pairs of lateral cusps, and often four. On the labial face of the crown the striae are 
coarse, sparse, and often bifurcated basally. They reach the tips of the lateral cusps 



BRITISH WEALDEN SHARKS 



297 




Fig. 6. Hybodus parvidens Smith Woodward. Teeth from the Wadhurst Clay of Hastings, 
Sussex, in labial view. a. P.11877, the holotype. b. P.11878 (paratype). c. P.11880 
(paratype) . 

and commonly reach the tip of the central cusp : in a sample of 37 teeth from Cliff 
End, the central cusp is striated to its tip in 22 ; it is striated to the tip in 10 of 13 
posterior teeth and in 12 of 24 anterior teeth. On the lingual face of the crown the 
ornament is very similar. It is interesting to note that in low-crowned posterior 
teeth, the striae on the lingual face of the crown tend to anastomose basally and 
form a reticular pattern (Text-fig. ys) approaching the pattern found in this region in 
low-crowned species like H. delabechei and H. brevicostatus (Text-fig. 10). At the 
base of the labial face of the central cusp a knob or accessory cusp is very commonly 
present. This accessory cusp is larger and more sharply defined in posterior teeth 
than in anterior (cf. Text-fig. 7A, b). In the Cliff End material this accessory cusp 
is present in more than half the teeth (in 28 of a sample of 40), and is commoner in 
posterior teeth (present in 13 of 16 posterior teeth and in 15 of 24 anterior teeth). 
The root in H. parvidens is typically hybodont, with foramina irregularly distributed 
on both surfaces, is moderately deep, and is turned lingually a little (Text-fig. 7). 
The above description is based on teeth from the Lower Wealden Ashdown Beds 
and Wadhurst Clay. Smith Woodward (1916 : 12) also recorded the species from 
the Weald Clay of Berwick, Sussex, but I have been unable to trace the specimen on 
which this record was based : it was probably an example of H. brevicostatus. H. 
parvidens also occurs in the Purbeck, as is shown by a sample of sixteen incomplete 
teeth from Friar Waddon, Dorset, and by 21349& and 21349^?, two teeth from the 
Middle Purbeck of Swanage, the first figured as H. ensis by Smith Woodward (1916, 
pi. 2, fig. 7). The picture presented by these Purbeck teeth (Text-fig. 8) is rather 
different from that in the Wealden material described above. The Purbeck teeth 
agree with the Wealden forms in size, but in most of them the crown is rather high, 
and where the ratio of crown height to tooth length is measurable (21349&, "^ it is 



GEOL. II, 7 



28 



BRITISH WEALDEN SHARKS 




2 mm 
Fig. 7. Hybodus parvidens Smith Woodward. An anterior (a) and a posterior (b) tooth in 
labial (i), lingual (2) and medial (3) view. P.46930-31, Ashdown Beds, Cliff End Bone-bed ; 
ClifiE End, Sussex. 



lower (I-5-I-7) than is usual in Wealden teeth. The striae on the labial face of 
the central cusp reach the tip in only four of the eighteen teeth : in six they cover 
between two-thirds and three-quarters of the cusp, in four about half the cusp, and 
in four less than half. Also, there is no accessory cusp at the base of the central 
cusp in any of the teeth, though there is an incipient cusp in two of them (Text-fig. 8b) . 
But in spite of these differences, I have little doubt in referring these teeth to H. 
parvidens : the examples with lower crowns exactly match those typical H. parvidens 
in which the accessory cusp is absent. 





2 mm 

Fig. 8. Hybodus parvidens Smith Woodward. Fragmentary teeth in labial view. a. 21349^, 
Purbeck ; Swanage, Dorset, b. P.46959, Upper Purbeck ; Friar Waddon, Dorset. 



BRITISH WEALDEN SHARKS 



299 



Among the material from the Paddockhurst bone-bed (Grinstead Clay) complete 
teeth are rare, but small central cusps and incomplete teeth (Text-fig. 9 a) are com- 
mon. Teeth of H. parvidens which agree with the typical forms from the Ashdown 
Beds and Wadhurst Clay occur, but are rather rare. In a sample of 37 teeth, four 
are typical low-crowned posterior teeth of H. parvidens, the other 33 are high- 
crowned forms. Of these 33 teeth, only three are striated to the tip of the central 
cusp (cf. 12 out of 24 in high-crowned teeth from Cliff End), and only three have an 
accessory cusp (cf . 15 out of 24 at Cliff End) . The striae in these teeth also tend to 
be finer and more nrmierous than in typical H. parvidens. 




1 mm 




Fig. 9. Hybodiis parvidens Smith Woodward. Teeth in labial view. 
Clay, Paddockhurst Bone-bed ; Paddockhurst Park, Sussex, b. 
Henfield, Sussex. 



A. P. 46944, Grinstead 
P.46971, Weald Clay ; 



Among the material from Tilgate Forest (Grinstead Clay) small teeth are very 
rare, but there are two examples of H. parvidens, 2850 and 3146. 

Among the material from the Weald Clay of Henfield I can find only two teeth 
which are referable to H. parvidens (Text-fig. qb) ; these are typical examples, very 
like teeth from the Grinstead Clay. Apart from these two specimens, all the high- 
crowned teeth from Henfield are of the type shown in Text-fig. 3, which are assigned 
to H. basanus for the reasons given on p. 292. The smaller low-crowned teeth (Text- 
fig. 13) are placed in H. brevicostatus for the reasons given on p. 306. H. parvidens 
is not known from any other locality in the Weald Clay or from any higher horizon. 

Affinities. The various samples of teeth of H. parvidens described above suggest 
that the history of the species was as follows. The species first appeared in the 



300 BRITISH WEALDEN SHARKS 

Middle Purbeck. Purbeckian teeth are rather high-crowned, with no accessory cusp 
and with rather short, tine striae. In the Lower Wealden (Ashdown Beds and Wad- 
hurst Clay) the species reached its typical and most distinctive form, with a crown 
which is of moderate height and has both striae to its tip and an accessory cusp in the 
majority of teeth. In the Grinstead Clay the species seems to trend towards the 
Purbeck form again, with an increase in crown height, a reduction in the height and 
an increase in the number of the striae on the central cusp, and a reduction in the 
incidence of the accessory cusp. Only two undoubted examples of H. parvidens 
are known above the Grinstead Clay. It seems very probable that the species did 
not become extinct, but evolved into H. basanus by a further increase in crown height 
and in the number of striae, coupled with an increase in size. The increase in size 
which marks the transition to H. basanus in the Weald Clay is perhaps correlated with 
the apparent extinction of the large-toothed H. ensis shortly before the Weald Clay 
was deposited. 

It is shown below (p. 308) that the Wealden (Ashdown Beds to Weald Clay) 
species H. hrevicostatus is also probably an offshoot of H. parvidens which originated 
near the base of the Wealden. 

Smith Woodward (1916 : 12) suggested a possible relationship between H. parvidens 
and the Upper Jurassic H. obtusits, a species in which the crown is moderately high, 
coarsely striated, and in which accessory cusps are very common. But the form of 
the earliest examples of H. parvidens in the Purbeck, where there is no accessory cusp 
and where the crown is higher and less strongly striated than it is in the typical 
forms, suggests that the species did not originate from H. obtusus but from some 
unknown small, high-crowned species in the Upper Jurassic. 

Hybodus hrevicostatus sp. nov. 
(PI. I, fig. 3 ; PL 2 ; PL 3, figs. 1-3 : Text-figs. 10-13) 

Diagnosis. Hybodus known by almost complete dentition and fin spines : 
nine paired files and a median symphysial file probably present in each jaw, 
dentition moderately heterodont ; teeth ranging (in one individual) from 8-o-i8-5 
mm. in length ; crown low and long, ratio of crown height (above the root /crown 
junction) to tooth length 3-25-6-25, crown not deeper than the root ; central cusp 
low, lateral cusps four or more, sometimes not recognizable in posterior and lateral 
teeth ; several accessory cusps often present on both labial and lingual margins of 
the crown, especially in the lower jaw ; crown with longitudinal occlusal crest and 
many coarse striae, often bifurcating basally, which reach the tips of the cusps and 
are interspersed in larger teeth with finer striae which fail to reach the tips of the 
cusps ; at the base of the lingual face of the crown the striae anastomose in a reticular 
pattern ; root deep, not turned lingually. Fin spines reaching about 18 cm. in 
length ; anterior edge keeled, lateral faces with 8-10 narrow, well spaced, discon- 
tinuous striae, posterior face with raised band bearing recurved denticles which lie 
in a single irregular series with occasional doubling, denticles extending proximally 
beyond the limit of the striae on the lateral surfaces. 



BRITISH WEALDEN SHARKS 301 

HoLOTYPE. BMNH no. P. 46973 (PL 2 ; PL 3, figs, i, 3), sixty-six teeth, fragments 
of calcified cartilage and an incomplete dorsal fin spine, the remains of a single in- 
dividual, from the Weald Clay of Henfield, Sussex. 

Material. In addition to the holotype, thirty isolated teeth and a fin spine. 

Horizons and localities. Ashdown Beds : Cliff End bone-bed, Sussex. Wad- 
hurst Clay : Hastings, Sussex. Grinstead Clay : Paddockhurst bone-bed, Sussex ; 
Tilgate Forest, Sussex. Weald Clay: Henfield, Sussex; Bookhurst, Surrey. Wealden 
Shales ; Atherfield Point and Cowleaze Chine, Isle of Wight. 

Description. The holotype was found in 1962 by Mr. J.F. Wyley immediately 
above the ' Pahidina ' limestone in the north-east corner of the Henfield pit. The 
teeth, calcified cartilage and fin spine were collected from an area of about two 
square feet : the bulk of the material was collected at one time but a few teeth have 
been found at the same point by Mr. Wyley in subsequent visits to the pit. There 
can be no doubt that this material represents the remains of a single individual. 
Since this is the first specimen of Hybodus in which it is possible accurately to re- 
construct the dentition with teeth free from matrix, and in which the upper and lower 
teeth can be distinguished, it will be described in some detail. 

I. The Dentition 

The teeth of the holotype were not found in natural association, but it has proved 
possible to reconstruct at least the anterior parts of the dentition with some accuracy. 
The teeth can be sorted into ' left ' and ' right ' (treating all the teeth as if they are 
from one jaw) by the asymmetry of the cusps, which point away from the symphysis 
in Hybodus, and by the asymmetry of the root. Many of the teeth can be matched 
with others to reconstruct successional files, and these files can be placed approxi- 
mately in their position in the mouth by the degree of asymmetry of the crown, and 
by the relative height and length of the teeth. When this had been done, of the 
fifty-seven more complete teeth four were median, twenty-three ' left ' and thirty 
' right ', and there appeared to be fourteen successional files, containing from two to 
eight teeth. Fourteen teeth could not be matched with any others. In these fourteen 
' series ', the measurements length of crown, length of root, depth of tooth, depth of 
crown, depth of root, breadth of crown and breadth of root were taken on each tooth 
(Table I), and the range of each measurement in each series was plotted against the 
inferred position in the mouth. The fourteen unmatched teeth were then measured 
and allocated their position in the mouth by reference to the plot. When this had 
been done it was clear that in each of the better represented tooth series there were 
two types of teeth : these represent the upper and lower jaws (see p. 303). 

General Features of the Dentition 

The dentition of H. brevicostatus (Text-fig. 10, PL 2) consists of a symphysial file 
of teeth in each jaw and probably nine paired files in each jaw. The largest teeth are 
the fifth paired file. In general features, the teeth are very like those of the Lower 



302 



BRITISH WEALDEN SHARKS 




5 mm 



Fig. io. Hybodus hvevicostatus sp. nov. Restoration of the dentition of the left side in labi 
VII and VIII in the upper jaw and III, VII and IX in the lower jaw are reversed drawinj 



Liassic H. delabechei Charlesworth (Smith Woodward 1889, pi. 10, figs. 1-5), with 
which they share a low crown, hardly exceeding the root in depth, a central cusp and 
four or more less conspicuous pairs of lateral cusps, and the ornament of very numer- 
ous prominent striae. The central cusp lies at or near the centre of the tooth in all 
but the most posterior files (Text-fig. 10, VIII and IX ; PL 2, figs. 5, 6), where it may 
be strongly eccentric or unrecognizable. The munber of lateral cusps is normally 
four on each side, but in the lateral and posterior teeth the lateral cusps may be 
more numerous or not recognizable. Accessory cusps occur very commonly on 
both the lingual and labial margins of the crown, particularly on the labial face of 
the mandibular teeth (see p. 305). 

There is a sharp occlusal crest running the length of the tooth from which strong 
striae pass to the base of the crown. These main striae often bifurcate basally, and 
between them there are weaker striae which fail to reach the occlusal crest. There 
is no significant difference between the strength of the striae on the labial and lingual 
faces of the crown : in some teeth they are stronger on the lingual face, in others on 
the labial. At the base of the lingual face of the crown the ridges anastomose in a 
reticular pattern. There is a good deal of variation in the ornament of the crown 
from one tooth to another, but this variation seems to have no significance between 
one jaw and the other or between tooth files. 

The roots of the teeth are typically hybodontoid (Casier 1947^ : 9). In all the 
teeth the root is moderately compressed, approximately equal to the crown both in 
depth and breadth, with a concave labial face and a flat or weakly convex lingual face. 
In all the teeth the root is without specialized foramina but has a large number of 
irregular foramina on both the lingual and labial faces. 

In histological structure (PL 3, fig. 3) the teeth are again typical of Hybodus (Jaekel 
1889, pi. 7, figs. 1,4). The crown is covered by a well marked and rather thick layer 
of enamel which contains fine fibrils in its basal part. Below the enamel there is a 



BRITISH WEALDEN SHARKS 
VI VII VIII 



303 



IX 




jview. ' s ', symphysial file, I-IX, paired files. All teeth drawn from the holotype ; V, VI, 
of teeth from the right side. P.46973, Weald Clay ; Henfield, Sussex. 

layer of very regular pallial dentine which makes up about one -fifth of the thickness 
of the crown. Below the pallial dentine the tooth consists of normal osteodentine. 
In the crown the osteodentine is dense, with many slender vascular canals : in the 
root it is much more spongy, with large, anastomosing vascular canals. 



Distinction between Upper and Lower Teeth 

As noted above, when the teeth of the holotype had been assigned to their positions 
in the jaws, among the better represented series (the symphysial and first five paired 
series) teeth of two types could be recognized. These clearly represent the upper and 
lower jaws. The upper and lower teeth can normally be distinguished by features of 
both the root and the crown, but the most reliable characters are in the root. In 
one type of tooth (Text-fig. 11 a) the labial face of the root is strongly concave, the 
base of the root is rounded, and there is normally a well marked furrow between root 
and crown on both labial and lingual faces of the tooth. In the second type of tooth 
(Text-fig. iib) the labial face of the root is less strongly concave, the base of the 
root is flattened in a well marked oblique shelf, and there is normally a very weak 
furrow at the junction of root and crown. The flattening of the base of the root is 





Fig. 



Hybodus brevicostatus sp. nov. Diagrammatic transverse sections of teeth from the 
lower (a) and upper (b) jaws, x 3. 



304 



BRITISH WEALDEN SHARKS 






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BRITISH WEALDEN SHARKS 305 

the most reliable method of distinguishing between the two types of teeth (PI. 2, 
cf. figs, la, 2a, and ^a, 4a). In the crown, those teeth with a rounded base to the 
root have significantly more accessory cusps on the margin of both labial and lingual 
faces. Taking only the teeth of the symphysial and first six paired files, accessory 
cusps are present on the labial face of all the teeth with a rounded base to the root. 
The number of accessory cusps on this face ranges from one to six, the mean being 
4*1. On the lingual face of these teeth accessory cusps are present in seventeen 
teeth out of twenty-six (65%), with a range of 1-3 and a mean of i-i. In teeth with 
a flattened base to the root, accessory cusps are present on the labial face in sixteen 
out of nineteen teeth (84%), with a range of 1-6 cusps and a mean of 1-7. On the 
lingual face of these teeth accessory cusps are present in six out of nineteen teeth 
(32%), with a range of 1-2 and a mean of o-6. The presence of an accessory cusp 
at the base of the labial face of the central cusp is particularly characteristic of teeth 
with a rounded base to the root, and there is very rarely an accessory cusp in this 
position in teeth with a flattened base to the root. This is reflected in the maximum 
breadth of the crown (Table I) which is the only dimension in which there is a 
significant difference between the two types of teeth. 

There is no absolutely reliable criterion by which these two types of teeth can be 
assigned to the upper or lower jaw, but comparison with associated dentitions of 
Hybodus obtusus from the Oxford Clay and with various species of Hybodus and 
Acrodus from the Lower Lias suggests that the teeth with the more numerous 
accessory cusps and the rounded base to the root are mandibular. The roots are 
visible in very few of the teeth in these associated dentitions, but comparison with 
isolated Jurassic teeth shows that the characters discussed above will serve to 
assign most teeth of Acrodus and low-crowned species of Hybodus to the appropriate 
jaw. 

Variation in Teeth with Position in the Jaws 

The main variations in the teeth from different parts of the jaws are clear from 
Text-fig. 10, PL 2 and Table I. The length and depth of both root and crown increase 
to a maximum in the middle of each jaw ramus, the length reaching a maximrun at 
the fifth paired file in the lower jaw and the sixth in the upper, the depth at the fourth 
file in the upper jaw and the fifth in the mandible. The depth of the root exceeds 
that of the crown in the symphysial, first, seventh, eighth and ninth files. The 
maximum breadth of the crown exceeds that of the root in all but the last two files 
in the mandible, while in the upper jaw the breadth of the root is always greater than 
that of the crown. 

1 
Other Material 

Teeth of H. brevicostatus first appear in the Ashdown Beds, and range through to 
the Wealden Shales, at the extreme top of the Wealden, but the species is rare through- 
out this time. 

Among the material from the Cliff End bone-bed (Ashdown Beds) there are six 
more or less complete teeth of the species, the best preserved of which is shown in 



3o6 



BRITISH WEALDEN SHARKS 





2 mm 



Fig. 12. Hybodus brevicostatus sp. nov. Tooth in labial (above) and lingual view. 
Ashdown Beds, Cliff End Bone-bed ; Cliff End, Sussex. 



P.iii 



Text-fig. 12. All these teeth are smaller than those of the holotype, with a maximum 
length of about lo mm., and they have fewer striae and fewer accessory cusps. 

In the Wadhurst Clay H. brevicostatus is represented by a fragment of a small 
tooth (P.11875) and by P.11876 (PL i, fig. 3), a large tooth, 15 mm. in length, 
probably a parasymphysial from the upper jaw, in which the crown is unusually high. 

In the Grinstead Clay there are thirteen teeth of H. brevicostatus among the material 
from the Paddockhurst bone-bed. These are all small (less than 9 mm. in length), 
low-crowned forms. 26027 from Tilgate Forest is a larger (11 mm.) example, again 
a low-crowned posterior tooth. 

Among the material from the Weald Clay of Henfield there is a number of teeth 
of H. brevicostatus in addition to the holotype. Most of these are normal teeth, as 
large as or a little smaller than those of the holotype, but two are very small examples 
(Text-fig. 13), P.46g84, less than 6 mm. in length and P. 46989, 4 mm. long. These 
small teeth agree with those of the holotype in shape and in the numerous accessory 
cusps, but in the few, coarse striae on the crown and in the clearly marked lateral 
cusps they resemble H. parvidens. One anterior tooth of H. brevicostatus (P.12812) is 
also known from the Weald Clay of Bookhurst, Surrey. 

The latest occurrence of H. brevicostatus teeth is P.13341, an incomplete para- 
symphysial tooth from the Wealden Shales of Atherfield Point, Isle of Wight. 



BRITISH WEALDEN SHARKS 



307 




2 mm 



Fig. 13. Hybodus brevicostatiis sp. nov. ? Juvenile teeth from the Weald Clay of Henfield, 
Sussex. A. P. 46984 in labial (above) and occlusal view. b. P. 46989 in labial view. 

2. Calcified Cartilage 

Together with the teeth of the holotype Mr. Wyley collected a number of fragments 
of heavily calcified cartilage. These include recognizable parts of the jaws and 
branchial arches, but they show nothing worthy of description. 



3. Fin Spines 

Found with the teeth and calcified cartilage of the holotype was a single incomplete 
fin spine (PL 3, fig. i). The British Museum (Natural History) contains one spine 
which agrees with this, P. 13268 from the Wealden Shales overlying the Hypsilophodon 
Bed, Cowleaze Chine, Isle of Wight (PL 3, fig. 2). These two spines agree almost 
exactly in size ; each must have had a total length of about 20 cm. Although they 
agree well in most characters, the angle of insertion of the Isle of Wight spine (in- 
ferred from the extent of the enamel ridges on the surface and the obliquity of the 
growth lines) was probably less than that of the Henfield spine. This suggests that 
the Isle of Wight specimen is an anterior fin spine and the Henfield is posterior, since 
in those hybodonts known by complete specimens the anterior fin spine lies more 
obliquely than the posterior (Brough 1935). 

The most striking feature of these fin spines, and the one which differentiates 
them from all other species of Hybodus, is that the enamel ridges on the distal part 
of the spine are very short, ending at or above the level of the lowermost denticles 
on the posterior face of the spine, and well above the apex of the groove on the 
posterior face of the proximal part of the spine which housed the basal cartilage of 
the fin. The trivial name of the species refers to these short ribs on the fin spine. 

The spines are not strongly arched, the anterior edge curving through about 35°, 
the posterior through 25°, and are strongly compressed. The anterior edge is 
keeled and there is a well marked angle between the lateral and posterior faces. 
On the lateral face the ridges of enamel are well spaced. In both specimens a median 



3o8 BRITISH WEALDEN SHARKS 

enamel ridge forms the keel on the anterior edge. In the Henfield spine (Pposterior) 
there are ten ridges on each side, five reaching the tip of the spine and five being 
confined to the proximal part. In the Isle of Wight spine there are eleven ridges 
on each side, seven of which extend to the tip. In the Henfield spine all the ridges 
end above the level of the lowermost denticle on the posterior face, the longest ribs 
being the first and second on the right side and the third on the left side. In the 
Isle of Wight spine the longest ridges are the first on each side, and they alone extend 
beyond the level of the lowest denticle on the posterior face. 

The denticles on the posterior face of the spine are in a single median series. This 
single series is clearly the result of unequal development of paired denticles, for 
occasionally the denticles are double (PL 3, figs, la, 2a), and in other cases at the base 
of a fully developed denticle there is the rudiment of a second. The side on which 
these rudimentary denticles occurs is variable, showing that the single series of 
denticles is not the result of suppression of either the left or right series of denticles, 
but of apparently random suppression of one of each pair. In the Isle of Wight spine, 
which is complete to the tip, there is a total of 27 denticles, of which two show doubl- 
ing and eight show rudiments of a second denticle, six on the right side and two on the 
left of a fully developed denticle. In the Henfield spine sixteen denticles are pre- 
served, one is doubled, and one has a rudiment on its left side. The denticles are 
very variable in shape : some are strongly hooked, either to the left or the right, 
some are straight, and while the majority are smooth, some, particularly towards 
the base of the spine, bear striae like those on the teeth. 

Lack of material at present prevents the preparation of thin sections of the spines, 
but broken surfaces show that in structure the spine agrees with H. aschersoni Stromer 
(1927 : 20, pi. 3, fig. 9, text-fig. 13) from the Cenomanian of Libya, in that in the 
exserted part of the spine the pulp cavity is surrounded by a zone of lamellar tissue 
which makes up about half the thickness of the wall of the spine, the outer part of 
the wall consisting of osteodentine. Passing back towards the base of the spine the 
zone of lamellar tissue becomes thinner until it disappears below the apex of the 
groove in the hind edge of the spine. Below this level the spine consists of osteo- 
dentine alone. 

Affinities. Teeth of H. brevicostatus from horizons below the Weald Clay suggest 
that in the history of the species between the Ashdown Beds and the Weald Clay 
there was an increase in size accompanied by an increase in the number of striae on 
the crown and in the number of accessory cusps. Small teeth from the Weald Clay 
of Henfield show that in the ontogeny of the species increase in size was accompanied 
by increase in the number of striae on the crown and reduction in the individuality 
of the lateral cusps. Both these facts suggest that the ancestor of H. brevicostatus 
was a rather small, low-crowned species, extant during or before the deposition of 
the Ashdown Beds, with sparse, coarse striae on the crown, three or four pairs of 
lateral cusps and a tendency to develop accessory cusps. These conditions are met 
by H. parvidens, and it seems very probable that H. brevicostatus is an offshoot of 
this species, probably originating near the base of the Wealden. 

The increase in ornamentation and reduction in the individuality of the lateral 



BRITISH WEALDEN SHARKS 309 

cusps which occux" with increased size is an interesting feature of this species. In 
the holotype it results in some of the teeth, especially the more posterior ones, having 
a very Acrodns-like crown, sometimes without distinguishable lateral cusps or with 
very numerous small ones. These conditions are just those found in some Upper 
Cretaceous species such as Synechodus illingworthi (Dixon) (Smith Woodward 
1911 : 220, pi. 46, figs. 5-7) from the Cenomanian zones of the English Chalk ; 
Hybodus brabanticus Leriche (1929 : 225, text-figs. 4, 5 ; 1930 : 105) and Acrodus 
dolloi Leriche (1929 : 228, text-figs. 6, 7) from the Lower Senonian of Belgium ; 
Hybodus grewingki Dalinkevicius (1935 : 256, pi. i, figs. 36-38) and Acrodus guedroyii 
Dalinkevicius (1935 : 256, pi. i, figs. 34, 35) from the Cenomanian of Lithuania ; 
Acrodus affmis Reuss (1845 : i, pi. 2, figs. 3, 4), Hybodus bronnii Reuss (1845 : 97, 
pi. 24, fig. 26, pi. 42, fig. 7) and Hybodus cristatus Reuss (1845 : 2, pi. 2, fig. 20), 
all from the Turonian Planerkalk of Bohemia ; and the teeth from the Upper Sono- 
nian of Aachen, Germany, described as Hybodus ? sp. and Acrodus ? sp. by Albers & 
Weiler (1964 : 4, 5, text-figs. 3, 8, 9, 42). These species are also the only marine 
hybodonts known by teeth in the Upper Cretaceous. The best known of them is 
Acrodus illingworthi Dixon, a species transferred to the heterodontid genus Synechodus 
by Smith Woodward (1891 ; 66). But there is no real evidence that this species is 
not a hybodont : all the teeth have typical hybodontoid roots, without the develop- 
ment of enlarged central foramina which is characteristic of Synechodus and other 
heterodontids (Casier 19476 : 2). The absence of hybodont fin spines in the English 
Chalk is not evidence against the species being hybodont, for only about twenty 
teeth are known, and judging by the conditions in H. brevicostatus an individual 
would produce well over two hundred teeth in its life but only two spines. It is also 
quite possible that these Upper Cretaceous hybodonts should have lost their fin 
spines. The syntypes and other material of A. illingworthi in the British Museum 
(Natural History) show a strong resemblance to H. brevicostatus, and with Leriche 
(1929 : 227) I believe the species should be named Hybodus illingworthi (Dixon). 
H. brabanticus Leriche is almost certainly a synonym of H. illingworthi , for Leriche 
included in his species some specimens figured by Smith Woodward (1911, pi. 46, 
fig. 7) as S. illingworthi, but these teeth (435 11) occur in association with other teeth 
which are undoubtedly H. illingworthi. There seems little to distinguish H. grewingki 
Dalinkevicius from H. illingworthi, and Dalinkevicius noted the resemblance between 
his material and Leriche's H. brabanticus. Reuss's species from the Turonian of 
Bohemia are large teeth of the same type as H. illingworthi and H. brevicostatus. 
The Upper Senonian Hybodus teeth described by Albers & Weiler are compared by 
them with H. illingworthi and H. brabanticus, which they closely resemble. Acrodus 
dolloi Leriche and A . guedroyii Dalinkevicius are based on large teeth which resemble 
Lower Lias species such as A. curtus {=^A. anningiae) and A. nobilis ; the Acrodus 
teeth described by Albers & Weiler are very like A . dolloi but smaller. There is no 
other evidence of Acrodus of this type in the Cretaceous, but these teeth, in size, 
shape and ornamentation, bear considerable resemblance to the large postero-lateral 
teeth of H. brevicostatus, and could be derived from this species by further increase 
in size, in the number of striae and in the loss of individuality of the lateral cusps. 



310 BRITISH WEALDEN SHARKS 

Fin spines of Hybodus 

Only in H. basanus and H. brevicostatus among Wealden and Purbeck sharks have 
fin spines been found in association with teeth. Smith Woodward (1916) suggested 
relationships between the various spines and teeth known from the British Wealden 
and Purbeck. His conclusions may be summarized as follows : 



Tooth 




Horizons 


Spine 


H. basanus 






Weald Clay 


? H. sulcatus Agassiz 


' teeth of the 


same 


general 






type as H. 


basanus ' 


Wadhurst Clay 


H. subcarinatus Agassiz 








Grinstead Clay 




H. ensis 






Middle Purbeck 


' almost identical with 
H. dorsalis Agassiz ' 


H. parvidens 






Wadhurst Clay 


p 


? 






Grinstead Clay 


H. striatulus Agassiz 


? 






Middle Purbeck 


H. strictus Agassiz 



Of the spines which Smith Woodward did not associate with species based on teeth, 
he compared H. striatulus with the Purbeck spines assigned to H. ensis, and H. strictus 
with H. subcarinatus and H. basanus. 

We now have more complete information on the ranges and relationships of the 
Wealden and Purbeck teeth of Hybodits. H. parvidens is a species which ranged 
from the Middle Purbeck to the Weald Clay, giving rise to H. basanus in the Weald 
Clay, the evolution of H. basanus involving an increase in size of the teeth. We 
should therefore expect the fin spines of H. parvidens to be similar to those of H. 
basanus but smaller, and to range from the Middle Purbeck to the Lower Weald Clay. 
These conditions are met by H. strictus Agassiz in the Purbeck (12-13 cm. long) and 
H. stibcarinatus Agassiz in the Wadhurst and Grinstead Clay (14-17 cm. long). 

The ' teeth of the same general type as H. basanus ' with which Smith Woodward 
associated H. subcarinatus Agassiz are shown above (p. 294) to belong in H. ensis : 
one would therefore expect them to be associated with spines of the same type as 
H. dorsalis Agassiz — this condition is met by H. striatulus Agassiz. 

A revised scheme of the (hypothetical) relationships between hybodont teeth and 
fin spines in the Wealden and Purbeck is as follows : 

Tooth Horizons Spine 

H. ensis Middle Purbeck- cf . H. dorsalis (Purbeck) 

Grinstead Clay H. striatulus (Wealden) 

H. parvidens Middle Purbeck- H. strictus (Purbeck) 

Weald Clay H. subcarinatus (Wealden) 

H. basanus Weald Clay ? H. sulcatus 

H. brevicostatus Ashdown Beds-Weald Clay H. brevicostatus 

In the Wealden and Purbeck there occur tuberculated fin spines which have been 
described as Asteracanthus {A. verrucosus Egerton 1854, Middle Purbeck, Swanage ; 
A. semiverrucosus Egerton 1854, Middle Purbeck, Swanage ; A. granulosus Egerton, 
1854, Wealden, Sussex). A spine from the Lower Neocomian of the Paris Basin is 
described as A. cf. acutus Agassiz by Leriche (1911 : 456, pi. 6, fig. i), and A . granulo- 
sus has also been recorded from the Lower Neocomian of Switzerland, where it occurs 



BRITISH WEALDEN SHARKS 311 

in the same beds as teeth of ' Strophodus ' {^ Aster acanthus) which according to 
Peyer (1946 : 54) are not derived fossils. But although Asteracanthus occurs in the 
marine Lower Cretaceous, it seems very unlikely that the British spines are true 
Asteracanthus — that is, that they are from a fish with ' Strophodus ' teeth, like the 
better known Jurassic species, for the large and conspicuous teeth could hardly have 
escaped notice, particularly since about a dozen spines of this type are known and in 
any individual there will be at least fifty teeth to each spine. The preservation of 
the spines is such that they can hardly be derived from older formations. The 
answer is probably that these spines are Hybodus in which the normal enamel ridges 
have broken up into separate tubercles as a stage in the reduction of the spine. 
This argument is supported by the presence of normal ridges on the distal (and first 
formed) parts of the spines, by the Wealden H. striatulus Agassiz (suggested above to 
belong to H. ensis) where the ridges are partially broken up into tubercles, and by 
the tendency in many Wealden Hybodus spines for the ridges to become fragmented 
proximally. 

Remarks on Hybodont Teeth 

This study has shown that where abundant isolated teeth of Hybodus from succes- 
sive horizons are available, the distinctions between species become very difficult 
to maintain. A great deal of parallelism and convergence is bound to occur in the 
evolution of sharks' teeth, and further uncertainty arises from the degree of hetero- 
donty of species (usually unknown) and from variability within a species. It is 
clear that the erection of new species of hybodont sharks on one or two isolated teeth 
is an extremely hazardous procedure. 

The changes which occur in the teeth of Wealden hybodont sharks from species to 
species and horizon to horizon allow one to draw some general conclusions about the 
correlation between various dental characters in hybodonts. These can be sum- 
marized as follows : 

{a) Characters which increase with increasing height of crown and decrease with 
decreasing height of crown. 

1. lingual curvature of crown 

2. compression of crown 

3. lingual curvature of root 

4. difference between ornament on labial and lingual faces of crown. 

{b) Characters which decrease with increasing height of crown and increase with 
decreasing height of crown. 

1. degree of heterodonty 

2. number of lateral cusps 

3. number and size of accessory cusps 

4. depth of root 

5. height of striae on crown 

6. bifurcation of and anastomosis between striae 



312 BRITISH WEALDEN SHARKS 

(c) Characters which increase with increasing size of tooth. 
I. number of striae on crown. 

This hst of characters showing correlation with the height of the crown or the size 
of the tooth includes almost all those commonly used to distinguish hybodont species. 
If all these characters are correlated with an obviously adaptive feature like crown 
height it is clear that it will often be impossible to distinguish on teeth alone between 
relationship, parallelism and convergence. There is therefore little hkelihood of any 
worthwhile subdivision of the genus Hybodus, now so large both in number of species 
and in range, on characters of the dentition. Jaekel's (i88g, 1898) attempt at a 
subdivision was largely based on features of the teeth which are obviously adaptive 
or are correlated with adaptive features : of the four genera which he proposed, only 
Polyacrodus, distinct on histological characters (thickness of pallial dentine), has 
been generally accepted. 

We are forced to look for other characters on which species of Hybodus may be 
grouped, but if the earliest (Lower Lias — H. delabechei etc.) and latest (Upper Wealden 
— H. basanus) species in which the anatomy is well known are compared, the only 
differences seem to be reduction of the cephaUc spines (present in males only) from 
two pairs to one, the appearance of symphysial teeth, and slight changes in the 
fin spines (increase in thickness of the lamellar tissue, reduction in the strength of 
the enamel ridges, reduction in the distance between the paired denticles on the 
hind edge, culminating in suppression of one of each pair in H. brevicostatus) . The 
first two of these differences are suitable for generic subdivision, but until their 
distribution is known in many more species no useful purpose is served by such 
division. 

Genus LONCHIDION Estes 1964 : 7 

Amended diagnosis. Small fresh-water hybodonts, weakly or strongly hetero- 
dont, known only by isolated teeth, cephalic spines and fin spines. Teeth low- 
crowned, elongated, crown not much deeper than root ; tricuspid anterior teeth 
present in some forms ; crown smooth or with sparse vertical striae, large labial 
projection or buttress below central cusp, lateral cusps absent or small and irregular ; 
root hybodontoid except in supposed tricuspid anterior teeth of advanced forms, 
where it is squatinoid ; pallial dentine of crown very thick, as in Polyacrodus. Fin 
spines with or without enamel ridges on lateral faces, with a single series of posterior 
denticles. Cephalic spines of normal hybodont type, without terminal barb. 

Type species. Lonchidion selachos Estes (1964 : 7, text-figs. 1-4), Lance Formation, 
Wyoming. 

Lonchidion breve sp. nov. 
(PL 5, fig. 3 ; Text-figs. 14-20, 29E) 
Diagnosis. Lonchidion known only by isolated teeth, 3-5 mm. or less in length ; 
no tricuspid anterior teeth, dentition weakly heterodont ; crowns of teeth rather 
short, with maximum breadth between half and one-quarter of the length ; crown 
smooth or very feebly striated, lateral and accessory cusps very small and irregular 
when present, labial process strong. 



BRITISH WEALDEN SHARKS 



313 



HoLOTYPE. P. 47024 (Text-fig. 14), complete tooth from the Paddockhurst bone- 
bed (Grinstead Clay) ; Sussex. 

Material. About one hundred and sixty teeth, mostly without roots. 

Horizons and localities. Ashdown Beds : Cliff End bone-bed, Chff End, 
Sussex. Wadhurst Clay : Telham bone-bed. Stone, Kent ; top of cliff near Hastings 
harbour, Sussex ; Ashiustwood, Sussex. Grinstead Clay : Paddockhurst bone-bed, 
Sussex. Weald Clay : Henfield, Sussex. Atherfield Clay : Perna Bed, Sandown 
Bay, Isle of Wight (probably derived from Wealden Shales — see p. 319). 

The teeth which are assigned to this species fall into three types which intergrade 
both morphologically and stratigraphically : they will be described separately as 
three subspecies. 




1 mm 

Fig. 14. Lonchidion hreve Ireve, sp. et ssp. nov. P. 47024, the holotype, a lateral tooth in labial 
(above), occlusal (centre) and lingual view. Grinstead Clay, Paddockhurst Bone-bed ; 
Paddockhurst Park, Sussex. 



Lonchidion breve breve sp. et ssp. nov. 
(PL 5, fig. 3 ; Text-figs. 14-16, 29E) 

Diagnosis. Lonchidion breve in which the crown of the tooth is normally smooth, 
without lateral or accessory cusps or striations. 

Holotype. P.47024, complete tooth (Text-fig. 14) from the Paddockhurst bone- 
bed. 

Material In addition to the holotype, about 120 teeth, mostly without roots. 

Horizons and localities. Ashdown Beds : Cliff End bone-bed. Wadhurst 
Clay: Telham bone-bed, Stone, Kent; Hastings, Sussex; Ashurstwood, Sussex. 



GEOL. II, 7 



29 



314 



BRITISH WEALDEN SHARKS 



Grinstead Clay : Paddockhurst bone-bed. Weald Clay : Henfield. Atherfield 
Clay : Perna Bed, Sandown Bay, Isle of Wight. 

Description. This subspecies, the typical and commonest form of the species, 
occurs throughout the Wealden. In the majority of specimens only the crown is 
preserved, and in many examples, especially those from the Cliff End and Paddock- 
hurst bone-beds, the teeth have undergone a good deal of post mortem abrasion. 
None of the teeth exceeds 3-5 mm. in length. The crown is normally completely 
smooth, without ornament or any indication of lateral cusps. There is a single, 
very low cusp which lies at or near the centre of the crown. The occlusal edge of 
the tooth is compressed into a more or less sharp ridge. Below the central cusp there 
is a strong rounded process or buttress on the labial face of the crown. The root 
plays no part in the formation of this process, and is overhung by it. A ridge nor- 
mally runs on to the labial process from the central cusp. On the lingual face of the 
crown there is a gentle swelling opposite the central cusp and the labial process. 
The occlusal surface of the crown is much longer than its base, so that the tooth 
is strongly waist ed at the junction of root and crown. 

The root is preserved in few specimens, and is always somewhat worn. It is 
typically hybodontoid (Casier 1947a : 9), with no specialized vascrdar foramina. On 
the lingual face of the root there is a row of vertically elongated foramina and on the 
labial face irregularly scattered foramina (Text-fig. 14). The root is almost as deep 
as the crown and only slightly shorter. 

In the type species of Lonchidion, the Late Cretaceous L. selachos Estes (see p. 330), 

A B 



0^ CS? 



^<^^^/ 










1 mm 

Fig. 15. Lonchidion breve breve sp. et ssp. nov. Three teeth from the Ashdown Beds, Qiff 
End Bone-bed ; Qiff End, Sussex, in labial (above), occlusal (centre) and lingual view. 
A. P.46993 ; B. P.46995 ; c. P.47005. 



BRITISH WEALDEN SHARKS 



315 



the histological structure of the crown is described (Estes 1964 : 8, text-fig. 2d) as a 
fan-shaped radiation of dentine tubules from one central cavity which originated 
along a central longitudinal axis. The tooth crowns of L. breve breve (PI. 5, fig. 3 ; 
Text-fig. 29E) agree with this description. Below a rather thick layer of enamel the 
crown is made up of pallial dentine containing long, subparallel, much branched 
tubules which arise from the tips of vascular canals which end shortly after entering 
the centre of the base of the crown. Estes compares this structure with that seen 
in teeth of the Triassic hybodont Palaeobates (Jaekel 1889, pi. 10, fig. 2 ; Seilacher 
1943, text-figs. 22, 29, 34), especially with P. nodosus Seilacher. But in Palaeobates 
(PI. 5, fig. 2) the dentine tubules are longer, parallel and more regularly branched 
than they are in Lonchidion, and they arise not from the tips of vascular canals 
but from a single open pulp cavity running the length of the tooth. The pallial 
dentine of Lonchidion is much more like that in the Triassic Polyacrodus, particularly 
in small species like P. minimus (Seilacher 1943, text-figs. 7-10 ; PL 5 fig. i), where 
there are just the same wavy, irregularly branched dentine tubules arising from 
short vascular canals near the base of the crown. In Polyacrodus the pallial dentine 
is a little thinner than it is in Lonchidion, perhaps because of the greater size of the 
teeth, but in other respects the correspondence is exact. 

The worn condition and absence of roots in most examples of this sub-species 
make it difficult to arrive at any clear idea of the variation in shape. The length 
of the crown ranges from 1-4-3 -4 mm., the maximum breadth of the crown (at the 
labial process) from 0-4-I-2 mm. In five complete teeth the depth ranges from 
1-0-I-35 mm., the ratio of depth to length from I-5-2-5. The crown is normally 
broader than deep, the ratio of maximum breadth to length ranging from I-9-3-8, 
and of maximum depth to length from 2 •7-4- 2. The shorter, deeper teeth, in which 
the labial process is large (Text -figs. 15A, i6a) are probably anterior ; longer teeth, 
still with a strong labial process (Text -fig. 15B), antero-lateral ; the longest teeth 



C^? 






1 mm 
Fig. 16. Lonchidion breve breve sp. et ssp. nov. Teeth in labial (above), occlusal (centre) and 
lingua] view, a, b. P.47324, P.47047 from the Weald Clay of Henfield, Sussex, c, d. 
P. 39015, P. 3901 1 from the Perna Bed, Atherfield Clay; Sandown Bay, Isle of Wight. 



3i6 



BRITISH WEALDEN SHARKS 



(Text-figs. 14, 15c) lateral, and shallow teeth with a weak labial process are probably 
posterior (Text-fig. i6b). 

At lower horizons there is no difficulty in distinguishing between L. breve breve 
and the other subspecies of the species, but at higher horizons, the Weald Clay of 
Henfield and the Perna Bed in the Atherfield Clay, examples occur in which there 
are weak accessory cusps (Text-fig. i6c) or lateral cusps (Text-fig. i6a, d). These 
specimens tend towards L. breve pustulatum, to which most of the Perna Bed teeth 
belong, and also towards L. selachos Estes, from the Upper Cretaceous of America 
(see p. 330). 

Lonchidion breve crenulatum sp. et ssp. nov. 
(Text-figs. 17, 18) 

Diagnosis. Lonchidion breve in which the occlusal margin of the tooth crown is 
irregularly crenulate, and in which the lingual and labial faces of the crown are 
weakly and sparsely striated. 

HoLOTYPE. P.47060 (Text-fig. 17B), tooth without root from the Paddockhurst 
bone-bed. 

Material. In addition to the holotype, twenty-five teeth, all without roots. 

Horizons and localities. Wadhurst Clay : Ashurstwood, Sussex. Grinstead 
Clay : Paddockhurst bone-bed. 












1 mm 



Fig. 17. Lonchidion breve crenulatum sp. et ssp. nov. Three teeth from the Grinstead Clay, 
Paddockhurst Bone-bed ; Paddockhurst Park, Sussex, in labial (above), occlusal (centre) 
and lingual view. a. P.47059. b. P.47060, the holotype. c. P.47066. 



BRITISH WEALDEN SHARKS 317 

Description. In this subspecies the form and proportions of the teeth do not 
differ significantly from those in L. breve breve. The largest tooth (Text-fig. 17c) is 
3-5 mm. long, similar in size to the largest L. breve breve. The occlusal crest of the 
teeth is produced into a number of small and irregular lateral cusps or beads. Both 
the Ungual and labial faces of the crown bear a number of weak, parallel, vertical 
striae, the striae being strongest at the central cusp. Both the beading of the occlusal 
crest and the striation of the crown are weakest in the short, deep anterior teeth 
(Text-fig. 17A), strongest in the longer, lower lateral and posterior teeth. 

Remarks. This subspecies is commonest in the Paddockhurst bone-bed, where it 
accounts for about half of the specimens of Lonchidion breve, the other half being 
L. breve breve. The fact that teeth of L. b. crenulatum from this horizon exhibit the 
same range of size and shape as those of L. b. breve (cf. Text -figs. 15, 17) and the ab- 




/ mm 

Fig. 18. Lonchidion breve crenulatum sp. et ssp. nov. P. 47081, a tooth from the Wadhurst 
Clay ; Ashurstwood, Sussex, in labial (above), occlusal (centre) and lingual view. 

sence of L. b. crenulatum at Cliff End and in the Telham bone-bed show that the two 
subspecies are distinct forms, not teeth from different parts of the mouth of the same 
fish. L. b. crenulatum is not known above the Grinstead Clay. In a sample of four 
teeth of Lonchidion from the Wadhurst Clay at Ashurstwood, Sussex, three are 
L. b. crenulatum (Text-fig. 18) and one is L. b. breve. L. b. crenidatum does not occur 
in the large sample of Lonchidion from the Cliff End bone-bed in the Ashdown Beds 
or the Telham bone-bed at the base of the Wadhurst Clay, nor is it known in the 
Purbeck, although the posterior teeth of L. heterodon (Text-fig. 25c) are very like 
it. In spite of this resemblance, it seems probable that L. b. crenulatum is a sub- 
species which evolved from L. b. breve, probably early in Wadhurst Clay time. 

Lonchidion breve pustulatum sp. et ssp. nov. 

(Text-fig. 19) 

Diagnosis. Lonchidion breve in which the occlusal crest of the crown is weakly, 
irregularly and finely crenulate, and in which the labial face of the crown bears a 
number of minute accessory cusps but no striations. 



3i8 



BRITISH WEALDEN SHARKS 



HoLOTYPE. P. 47085, (Text-fig. 19B), tooth without root from the Perna Bed, 
Atherfield Clay, Sandown Bay, Isle of Wight (? derived from the Wealden Shales — see 
below) . 

Material. In addition to the holotype, fourteen teeth, all without roots. 

Horizon and locality. Perna Bed, Atherfield Clay : Sandown Bay, Isle of 
Wight. 

Description. In this subspecies the form and proportions are again very similar 
to those in L. breve breve. The teeth range in length from I-8-3-6 mm., in breadth 
from 0-6-I-3 mm., and in the ratio of breadth to length from 2-0-3-6. The occlusal 











/ mm 

Fig. 19. Lonchidion breve pustulatum sp. et ssp. nov. Teeth from the Perna Bed, Atherfield 
Clay ; Sandown Bay, Isle of Wight, in labial (above), occlusal (centre) and lingual view. 
A. P.47088. B. P. 47085, the holotype. c. P. 47089. 

crest of the teeth is produced into irregular lateral cusps or beads, and these are 
more numerous than they are in L. breve crenulatum. The labial and lingual faces 
of the crown are without striae. The labial process, below the central cusp, is not so 
well marked off from the body of the crown as it is in the other subspecies (Text-fig. 
19B, c), and the ridge on its occlusal surface bears one or sometimes two small cusps. 
Level with the more labially placed of these cusps on the labial process there is a 
series of very small accessory cusps on the labial surface of the crown, which extends 
as a shallow shelf at this level. The strength and number of these accessory cusps 
seem to be correlated with the breadth of the tooth (Text -fig. 19), but not with its 
length, and no real distinction between anterior and posterior or lateral teeth can 
be made. 



BRITISH WEALDEN SHARKS 



319 



Remarks. This subspecies is known only in the Perna Bed, at the base of the 
Lower Greensand. Most of the fish teeth from this horizon are derived from the 
underlying Wealden Shales (Casey 1961 : 505), and this is probably true of the speci- 
mens of Lonchidion, a genus otherwise unknown in marine beds. The teeth are 
not much rolled or waterworn, but their small size will probably account for this. 
Together with L. b. pustulatum in the Perna Bed there occur teeth which are referred 
to L. b. breve but which show faint traces of the lateral and accessory cusps which 
characterize L. b. pustulatum (Text-fig. i6c, d) : Similar examples also occur at 
Henfield (Text-fig. i6a). These forms leave little doubt that L. b. pustulatum has 
evolved directly from L. b. breve by extension of the labial face of the crown into a 
shelf bearing accessory cusps. Teeth of L. b. pustulatum are very like the smallest 
posterior teeth of the ptychodont genus Hylaeobatis, suggesting a possible relationship 
which is discussed on p. 341. 

The Arrangement of the Teeth 

Many of the teeth of Lonchidion breve have a strong wear facet on the occlusal 
margin of the crown, the occlusal crest and central cusp being planed off into a level 
surface. This indicates, as one would deduce from the form of the teeth, that L. 
breve was a durophagous form, grinding its food by rubbing the upper and lower 
teeth across one another. In many specimens of L. breve there is also a vertical 
groove or scar in the centre of the lingual face of the crown : this is a pressure scar 
formed by the tip of the labial process of the succeeding tooth, which shows that 
successive teeth in each file were in contact or nearly so. If the tip of the labial 
process of each tooth touched the centre of the lingual face of the crown of the preced- 
ing tooth, there would be a space between the tapering lateral parts of the teeth — 
such spaces are disadvantageous in durophagous forms, and they must have been 
filled by the lateral parts of the teeth of the adjacent file (Text-fig. 20). It follows 
that in L. breve there was overlap between the files of teeth, and that there must 
therefore have been regular alternation between the teeth in adjacent files : this is 
an advanced character recalling the rays, which does not normally occur in hybodonts 




Fig. 20. Lonchidion breve sp. nov. Diagram showing the probable arrangement of adjacent 
files of teeth in occlusal view, x 15. Based on the holot3rpe (Text-fig. 14). 



320 



BRITISH WEALDEN SHARKS 



and heterodontids (Casier 1953 : 40), although it is present in the Triassic fresh- water 
form Lissodus africanus (Brough 1935, pi. 2, figs. 2, 3), at least in the anterior and 
lateral parts of the jaw. 

Lonchidion striatum sp. nov. 
(Text-figs. 21, 22) 

Diagnosis. Lonchidion known only by isolated teeth, 4-5 mm. or less in length ; 
no tricuspid anterior teeth, dentition weakly heterodont ; crowns of teeth elongated, 
maximum breadth rarely less than one-quarter of the length ; occlusal crest of teeth 
strong, lateral cusps very weak and irregular ; both labial and lingual faces of 
crown with strong vertical striae, occasionally bifurcated basally, which diverge 
from the occlusal crest but do not reach the base of the crown ; labial process small 
and weak. 

HoLOTYPE. P. 47103 (Text-fig. 21c), a complete tooth from the Weald Clay of 
Henfield, Sussex. 

Material. In addition to the holotype, one hundred teeth, of which nine are 
complete. 

Horizon and locality. Weald Clay : Henfield, Sussex. 

Description. The teeth of this species from Henfield are all well preserved, with 
little or no wear, but the root is present in only nine of the one hundred examples. 
The teeth range in length from i-i to 4-2 mm. In shape and proportions the teeth 
are similar to those of L. breve breve and L. b. crenulatum except that they are more 
elongated. Only in two teeth out of fifty does the ratio of maximum breadth to 
length of the crown fall below 4-0, the mean ratio being 5-9 (compared with about 
2-7 in L. b. breve). The longitudinal occlusal crest of the crown is strongly marked, 




D 



xo^v^\rn^^^\^\^i'^^TTT^ 



' mm 



Fig. 21. Lonchidion striatum sp. nov. Teeth from the Weald Clay of Henfield, Sussex, in 
labial (above), occlusal (centre) and lingual view. a. P. 47094. b. P. 47095. c. P. 47103, 
the holotype. d. P. 47 106. 



BRITISH WEALDEN SHARKS 



321 



and there is a low central cusp. The occlusal crest is roughened and jagged, as in 
L. b. cremilatum, but distinct lateral cusps are not recognizable. Diverging from the 
occlusal crest there are well marked vertical striae on both labial and lingual faces 
of the crown, the striae being stronger and more numerous than in L. b. crenulatum. 
The striae are occasionally bifurcated basally, especially at the central cusp, where 
they are longest and strongest. The striae do not reach the base of the crown. 
The labial process is weaker than in L. breve, particularly in the more elongated 
teeth, where it may almost disappear (Text-fig. 21A). The striae are weakest in the 
smaller and shorter teeth (Text-fig. 22a). 

The root is about as deep as the crown, and is hybodontoid, without specialized 
foramina, just as in L. breve. In histological structure the crown agrees with that of 
L. breve in being made up almost entirely of pallial dentine. 





^^^^^^^ 




/ mm 

Fig. 22. Lonchidion striatum sp. nov. Teeth from the Weald Clay of Henfield, Sussex, in 
labial (above), occlusal (centre) and lingual view. a. P. 47130. b. P. 47138. 

There is little variation in the shape of the teeth, and the dentition must have 
been weakly heterodont. The shorter, broader teeth, with the weakest striae 
(Text-figs. 21B, 22a), are probably anterior, longer teeth (Text -figs. 21c, 22b) lateral, 
and long, slender teeth, with a very small labial process (Text-fig. 21A, d), posterior, 
similar variations being evident in L. breve crenulatum (Text-fig. 17). 

Affinities. Lonchidion striatum differs from L. breve crenulatum only in the 
greater length of the teeth (ratio of breadth to length usually well over 4-0, always 
less than 4-0 in L. b. crenulatum), stronger and more numerous striae, and weaker 
labial process. There can be little doubt that L. striatum, known only in the Weald 
Clay, has evolved directly from L. breve crenulatum, which is unknown above the 
Grinstead Clay, by elongation of the teeth and strengthening of the surface ornament. 
The reduction of the labial process in the lateral and posterior teeth of L. striatum 



322 



BRITISH WEALDEN SHARKS 



means that there would be httle space between the lateral parts of successive teeth 
(cf. Text -fig. 20) and little or no overlap between adjacent files of teeth, producing 
a more Hybodus-like dentition than in L. breve. 

In shape, teeth of L. striatum are similar to some of the Triassic species of Acrodus 
and Polyacrodus such as the Spitzbergen species Polyacrodus pyramidalis Stensio 
(1921, pi. 2, figs. 21-26), Acrodus spitzbergensis Hulke (Stensio 1921, pi. 2, figs. 1-19), 
A. vermiformis Stensio (1921, pi. 2, figs. 20, 21) and A. oppenheimeri Stensio (1921, 
pi. 3, figs, i-ii) though the teeth of all these species are larger than those of L. 
striatum. 

Lonchidion rhizion sp. nov. 

(Text-figs. 23, 24) 

Diagnosis. Lonchidion known only by isolated teeth less than 2 mm. in length ; 
no tricuspid anterior teeth, dentition strongly heterodont ; crowns of teeth very 
short and broad, maximum breadth more than half of the length ; occlusal surface 
of teeth formed entirely by labial surface of crown and of labial process, which is 
much enlarged and probably overlapped the crown of preceding tooth ; crown 
smooth, without ornament or lateral cusps ; lingual surface of crown with pair of 
depressions or sockets separated by central crest which probably articulated with 
pulp cavity of succeeding tooth ; pulp cavity small, root absent or very small. 

HoLOTYPE. P. 47144, tooth without root (Text -fig. 23B), Cliff End bone-bed, 
Cliff End, Sussex. 

Material. In addition to the holotype, thirty-five teeth, all without roots. 

Horizons and localities. Ashdown Beds : Cliff End bone-bed. Cliff End, 
Sussex, (25 teeth). Wadhurst Clay: Telham bone-bed, Stone, Kent (5 teeth). 
Grinstead Clay : Paddockhurst bone-bed, Paddockhurst Park, Sussex (5 teeth). 

Description. The teeth assigned to this species vary from examples which do 
not differ much from L. breve breve (Text -fig. 23A) to extremely specialized types which 




1 mm 



Fig. 23. Lonchidion rhizion sp. nov. Teeth from the Ashdown Beds, Chff End Bone-bed ; Cliff 
End, Sussex, in Ungual (i), labial (2), medial (3), occlusal (4) and basal (5) view. a. ? anterior 
tooth P.47143. B. ? lateral tooth, P. 47144, the holotype. c. ? posterior tooth, P.47150. 



BRITISH WEALDEN SHARKS 323 

would hardly be interpreted as sharks' teeth were they not linked by intermediate 
forms with the more recognizable specimens. All the teeth are very small, ranging 
in length from i-o to 1-9 mm., and they are broad and deep ; the ratio of maximum 
breadth to length ranges from 0-75 to 2-0, that of depth to length from i-o to 2-4. 

The deeper, narrower teeth (Text-fig. 23A) are similar to the (presumed) anterior 
teeth of L. breve breve (Text-fig. 15A). In these teeth the crown is almost as deep as 
long, and is deeper than broad. The crown is pentagonal in lingual and labial view, 
two sides of the pentagon forming the occlusal crest and meeting in a rounded central 
cusp, two sides sloping inwards towards the base, and one side forming the base. 
The lingual face of the crown is almost smooth, with a slight swelling below the central 
cusp and a shallow depression on each side of this. In occlusal view the crown is 
triangular, the longest side being the occlusal crest and the apex the labial process. 
The labial process is much broader than it is in L. breve, and its flat upper surface 
merges with the upper part of the labial face of the crown to form the smooth, 
sloping occlusal surface of the tooth. There is no ridge running on to the labial 
process from the central cusp as there is in L. breve. The underside of the labial 
process is hollowed out, this hollow being continuous basally with the small pulp 
cavity of the crown. 

More specialized, but linked by intermediate forms with the teeth described above, 
are teeth of the type shown in Text-fig. 23B. In these the crown is shallower but 
broader, and is almost as broad as long. The lingual face of these forms bears a 
central vertical crest with a well marked, circular depression on each side of it. The 
occlusal margin of the crown is lower and more gently rounded, the central cusp 
hardly recognizable in labial or lingual view but showing in occlusal view as a knob 
at the top of the central crest on the lingual surface. The labial process is longer than 
in the forms described above, and the occlusal surface formed by its upper surface 
is much larger. In medial view, the labial process curves labially and basally well 
beyond the rest of the crown. The pulp cavity in the basal surface of the crown is 
very small, and extends only into the proximal part of the labial process. 

The most highly specialized teeth (Text-fig. 23c) have the crown shallower, the 
occlusal margin flatter and more gently rounded, and the crest and paired depressions 
on the lingual surface more strongly marked. In these forms the depressions on the 
lingual face are in the form of rounded sockets. The labial process is enormously 
enlarged, so that the breadth of the tooth is greater than its length (ratio of breadth 
to length as low as 0-75). The process projects labially in a long, tongue-like flange. 
The pulp cavity in the basal surface of the crown is very small, and hardly extends 
into the base of the labial process. 

In none of the teeth of this species is any trace of a root preserved. The basal 
surface of the crown and the wall of the small pulp cavity are always quite smooth 
and without the foramina usually present in teeth where the root has been broken 
or abraded off. In the more specialized teeth the pulp cavity and basal surface of 
the crown are both so small that if a root were present its value as an anchor would be 
neghgible. The form of the teeth and of the basal surface and pulp cavity suggest 
that there may have been no root in this species, especially in the shallower, more 



324 



BRITISH WEALDEN SHARKS 



specialized teeth. This conclusion is supported by the inferred mode of articulation 
of the teeth (see below), which seems to allow no room for a root. 

The Arrangement of the Teeth 

As shown in Text-fig. 23, there is a great deal of variation in the teeth of L. rhizion, 
involving the depth of the tooth, the length of the labial process, and the symmetry 
of the tooth. I can find no correlation between these characters, except that the 



B 








Fig. 24. Lonchidion rhizion sp. nov. Diagram showing possible modes of arrangement of 
the teeth, X15 approx. For explanation see text. Based on the holotype (Text-fig. 236). 
A, B, C3 in medial view ; c^ in basal view ; Cj in occlusal view. 

labial process is normally longer in shallower teeth, but deep, asymmetrical teeth 
with rather long labial processes do occur. As yet, there is therefore no means of 
assigning the different teeth to a position in the mouth, but it is clear that the dentition 
of L. rhizion must have been strongly heterodont. 

In none of the teeth of L. rhizion is there any well marked wear facet on the occlusal 
surface of the tooth. In L. breve there is often a vertical pressure scar in the centre 
of the lingual surface of the crown, caused by the tip of the labial process of the 
succeeding tooth : in L. rhizion there is never such a scar, but instead there is a 
central crest and a pair of depressions. These depressions are very weak in the 
deepest teeth with the largest pulp cavity (Text-fig. 23A), but become stronger as the 
depth of the tooth and the size of the pulp cavity decrease, until in the shallowest 
teeth (Text-fig. 23c) they are deep sockets. The apparent correlation (Text-fig. 23) 
between length of the labial process and strength of the lingual depressions does not 
hold good for all specimens. The shape and position of these Ungual depressions 
and the crest between them suggest that they served for articulation between the 
teeth, and this is supported by the fact that they are strongest in teeth in which the 
pulp cavity (and basal surface of attachment) is smallest. 



BRITISH WEALDEN SHARKS 325 

There are three possible ways in which the successive teeth in L. rhizion might 
have been arranged. 

1. As in L. breve (Text-fig. 20), the teeth might have been arranged in a simple 
file, but without contact between successive teeth (since there is never a pressure 
scar on the lingual face which might be caused by the tip of the labial process of the 
succeeding tooth). This scheme is shown in Text-fig. 24A. It is very unhkely that 
the teeth were arranged in this way because it allows no function for the paired 
depressions and central crest on the lingual face of the crown of the shallower teeth ; 
because the enlargement of the labial process in the shallower teeth would also be 
without apparent function, and because the teeth with very long, slender labial 
processes (Text-fig. 23c) would give an unsatisfactory dental series. 

2. The teeth might have been arranged as in Text-fig. 24B, with the labial surfaces 
forming the occlusal surface and the tip of the labial process of each tooth ending at 
the central cusp of its predecessor. Although this scheme gives a satisfactorily 
smooth occlusal surface to the dental series, it is open to the same objections as the 
scheme outlined above. The only function for the paired depressions on the lingual 
surface of the crown in this scheme would be to receive a pair of processes from the 
root of the succeeding tooth : this is unlikely, for if the roots were large enough to 
abut against the preceding tooth it is probable that they would be preserved in some 
specimens. 

3. The teeth might have been arranged as in Text-fig. 24c, with the labial process 
of each tooth overlapping the central cusp of its predecessor. Manipulation of 
plasticine models of the teeth shows that in this type of arrangement the boss at 
top of the crest in the centre of the lingual surface would have fitted in the labial part 
of the pulp cavity of the succeeding tooth, and that the paired depressions would 
have articulated with the ridge or knob on each side of the pulp cavity at the base 
of the labial process. In this arrangement the radius of the tooth whorl would have 
been rather small, since a satisfactory fit between successive teeth is only obtained if 
their axes differ by about forty degrees. This scheme gives a function to the crest 
and paired depressions on the lingual face of the crown, gives a functional explanation 
of the elongation of the labial process, and gives each tooth series a radula-Uke 
occlusal surface which should be highly efficient. It does not give an entirely 
satisfactory fit between either the deep teeth with a short labial process (Text-fig. 23A) 
or the shallow teeth with a very long labial process (Text-fig. 23c), but with most of 
the specimens it is satisfactory. If the teeth were arranged in this way, part of the 
pulp cavity was occupied by the lingual crest of the preceding tooth and not by a 
root. In the shallowest teeth with the longest labial process the pulp cavity is so 
small that the articulation between the teeth would have left only a narrow channel 
linguaUy, through which nerves and vessels could have passed into the pulp cavity. 

It is suggested, pending the discovery of more complete material, that Lonchidion 
rhizion was a shark in which a flattened but serrated grinding dentition was evolved 
by the labial process of each tooth overlapping the crown of its predecessor ; that this 
overlapping entailed drastic reduction and eventual loss of the root, and that this 



326 



BRITISH WEALDEN SHARKS 



was compensated for by the development of complex interlocking articulations be- 
tween successive teeth. 

Affinities. Lonchidion rhizion is known only from the Ashdown Beds (Cliff End 
bone-bed), where it is not common, and from the Wadhurst Clay (Telham bone-bed) 
and Grinstead Clay (Paddockhurst bone-bed), where it is rare. The least speciaUzed 
teeth are very like the (presumed) anterior teeth of L. breve breve, and it is probable 
that the species evolved from L. breve, primarily by flattening and broadening of the 
labial process. L. rhizion is the only known shark in which some, at least, of the 
teeth were without roots. 



Lonchidion heterodon sp. nov. 

(Text-fig. 25) 

Diagnosis. Lonchidion known only by isolated teeth, less than 4-0 mm. in length ; 
tricuspid symphysial teeth probably present, dentition heterodont ; crowns of 
teeth moderately elongated, ratio of breadth to length 2-5-4-5 (except in the presumed 
symphysial teeth, where the ratio is less than 2-0), crown shallower than broad in 
lateral teeth, deeper than broad in anterior and posterior teeth ; teeth Hybodus-Vik.e, 
with low central cusp and three pairs of lateral cusps (one pair in symphysial teeth), 
with striae diverging from occlusal crest, striae few and weak in anterior and posterior 
teeth, numerous and strong in large lateral teeth ; labial process strong in anterior 
and posterior teeth, weak in lateral teeth. 

HoLOTYPE. P.47188 (Text-fig. 25A), a lateral tooth without root from the Upper 
Purbeck, Friar Waddon, Dorset. 

Material. Twelve teeth, all without roots. 

Horizons and localities. Upper Purbeck : Friar Waddon, Dorset (6 teeth). 
Ashdown Beds : Cliff End bone-bed. Cliff End, Sussex (5 teeth). Wadhurst Clay : 
Hastings, Sussex (i tooth). 




Fig. 25. Lonchidion heterodon sp. nov. Teeth in labial (above) and occlusal view, c also in 
lingual view (below). A. P.47188, lateral tooth, the holotype. b. P.47189, antero-lateral 
tooth, c. P.47192, posterior tooth, d. P. 47197, symphysial tooth tentatively referred to 
this species, a, b, c from the Upper Purbeck ; Friar Waddon, Dorset ; d from the Ashdown 
Beds, Cliff End Bone-bed ; Cliff End, Sussex. 



BRITISH WEALDEN SHARKS 327 

Description. The holotype is a broad, low-crowned tooth which is Hybodus-like 
in appearance, with a low central cusp and three pairs of lateral cusps. The occlusal 
crest is well marked. Both labial and lingual faces of the crown bear strong striae 
which reach the occlusal crest at the central and lateral cusps but fail to do so between 
the cusps. The striae are occasionally bifurcated basally, and are stronger and longer 
on the labial face than on the lingual. The labial process, below the central cusp, is 
poorly marked. Both the basal surface of the crown and the lingual surface are 
concave, and the tooth is strongly asymmetrical. The material contains seven teeth 
of this type, ranging in length from 2-5 to 4-0 mm., and in the ratio of breadth to 
length from 3-0-4-5. The asymmetry and curvature of the basal surface and occlusal 
crest are particularly characteristic of these teeth. Smaller teeth of this type grade 
into the type shown in Text -fig. 25B. which are about 2 mm. in length and have the 
crown higher and narrower. In these teeth the central cusp is stronger and the 
lateral cusps weaker than in the larger teeth, while the labial process is larger and the 
striae fewer and shorter. These teeth are still weakly asymmetrical, and have the 
basal surface and the labial surface concave. In the small tooth shown in Text-fig. 
25c these trends go further : the crown is higher and narrower, the central cusp and 
labial process are accentuated, the lateral cusps and striae reduced. There is little 
difference between this tooth and examples of Lonchidion breve crenulatum (Text-fig. 
17) from the Grinstead Clay. 

Tentatively included in this species is the tooth shown in Text -fig. 25D. This is 
bilaterally symmetrical and must be a symphysial tooth. There is a large central 
cusp, a single pair of sharp lateral cusps, a large labial process, and a few coarse 
striae. In this tooth (length 1-7 mm.) the breadth and depth are almost equal, and 
the ratio of breadth to length is about 1-7, much less than in the other teeth of the 
species. 

The material of this species, though very limited, suggests that the dentition was 
strongly heterodont and consisted of small, high-crowned symphysial teeth (Text-fig. 
25D), large, broad, low-crowned lateral teeth (Text-fig. 25 a), smaller, narrower 
antero-lateral and postero-lateral teeth (Text-fig. 25B), and posterior teeth which are 
very like Lonchidion breve crenulatum (Text-fig. 25c). 

The root is not preserved in any specimen. In histological structure (seen in 
peels taken from ground, etched surfaces of a fragment of a lateral tooth from the 
Upper Purbeck) the crown of this species agrees exactly with Lonchidion : the 
crown is made up entirely of pallial dentine, the long, branched dentine tubules 
radiating from vascular canals at the base of the crown. 

Affinities. This species, though poorly known, is of interest and importance as 
the only record of Lonchidion in the Jurassic. The smallest teeth of the species, 
presumably posterior, are almost identical with those of L. breve crenulahim from the 
Grinstead Clay, but are linked by a series of intermediate forms with teeth like the 
holotype (Text-fig. 25A) which are very Hybodns-Yike (cf . H. parvidens — Text-fig. 7B ; 
H. brevicostatus — ^Text-fig. 13B) though still retaining the very thick pallial dentine 
which is typical of Lonchidion. L. heterodon is rare at Cliff End, in the Ashdown 
Beds ; and is known above this only by a single lateral tooth from the Wadhurst 



328 



BRITISH WEALDEN SHARKS 



Clay. If we assume that L. heterodon, as the earliest known species of Lonchidion 
shows the most primitive condition, the genus must have originated as a heterodont, 
low-crowned, Hybodus-like form with very thick palhal dentine. The more homodont 
species, L. breve, would then have evolved by adoption in all the teeth of the slender, 
high-crowned, almost smooth form of the posterior teeth in L. heterodon, and by 
increasing the overlap between adjacent files of teeth. 

Fin Spines and Cephalic Spines of Lonchidion 

In the late Cretaceous species Lonchidion selachos, Estes (1964 : 9) was able to 
include fragmentary fin spines and cephalic spines from locaUties at which teeth of 
the species occur. There is little doubt of the association between the teeth and the 
spines since L. selachos is the only hybodont known from the deposits. In the British 
Wealden and Purbeck the matter is complicated by the presence of small and im- 
mature hybodonts of other species. 



i,|/ 





B 




Fig. 26. Lonchidion sp. a. Dorsal fin spine, a restoration based mainly on P.29995, Wealden ; 
Isle of Wight, P.12815, Weald Clay ; Bookhurst, Surrey, and P.47208, Weald Clay ; Henfield, 
Sussex, in lateral (left) and posterior view, x li approx. The broken line marks the plane 
of the section shown in PI. 3, fig. 4. b. CephaUc spine, P. 1 1895 from the Wadhurst Clay; 
Brede, Hastings, Sussex, in dorsal (below) and lateral view, X3-5 approx. c. Incomplete 
cephalic spine in lateral view, P. 47207, Weald Clay ; Henfield, Sussex, x 6-5. 



BRITISH WEALDEN SHARKS 329 

In L. selachos the cephalic spine (Text-fig. 27E) has a weakly lobed root, no barb 
at the tip, and is ornamented with small tubercles which are drawn out into ridges 
distally and are pointed anteriorly. The spine is about lo mm. in length, roughly 
2-5 times as long as the average lateral tooth of the species. Among the material 
from the Cliff End bone-bed there are several worn and incomplete cephalic spines 
of small size : a complete spine of this type from the Wadhurst Clay was figured by 
Smith Woodward (1916, pi. i, fig. 4). These spines (Text-fig. 26B) differ from the 
cephalic spine assigned to L. selachos in that the root is much wider and more sharply 
divided into three lobes, and the enamelled exserted part of the spine lies at a much 
shallower angle to the root and does not curve back so far. In all the specimens from 
Cliff End the enamelled part of the spine is perfectly smooth, but this could well be 
due to abrasion, for in a similar spine (P.12813) from the Weald Clay of Bookhurst, 
Cranley, Surrey, there are vertical striae at the base (the only part preserved) of 
the exserted portion. These cephalic spines are tentatively referred to Lonchidion : 
probably they belong to L. breve, the commonest species at Cliff End. 

A single cephalic spine from the Weald Clay of Henfield (P. 47207, Text-fig. 26c) is 
also probably a Lonchidion. The root is almost entirely missing but the crown curves 
back in just the same way as in L. selachos, much more strongly than in the Cliff End 
and Wadhurst Clay spines. The tip of the spine is smooth and without a barb, but 
the proximal part of the crown is ornamented with about twelve sub-parallel striae, 
quite a different form of ornamentation from L. selachos. This cephalic spine is 
perhaps from L. striatum, the dominant species of Lonchidion at Henfield. 

The fin spines of L. selachos (Text-fig. 27F) are all incomplete, so that the total 
length cannot be measured, but they reached at least 5 mm. in breadth, and were 
probably about 5 cm. long. The surface is apparently without enamel ridges and is 
marked only by the weak striations of the texture of the superficial osteodentine. 
There is a single series of denticles on the posterior face. 

There are no complete fin spines among the bone-bed material described here, and 
the fragments of spines from Cliff End and Paddockhurst are too worn for description. 
But fragmentary spines from the Weald Clay of Henfield and Bookhurst, Cranley, 
Surrey, and from an unknown horizon (? Perna Bed) in the Isle of Wight seem to be 
referable to Lonchidion, although it is only in the few examples where part of the 
inserted base is preserved that one can distinguish with certainty between small 
spines and the tips of large spines of Hybodus. From these fragments one can recon- 
struct the fin spine of Wealden Lonchidion (Text-fig. 26a). The spines reached about 
7 cm. in length and were almost straight, the posterior border of the spine showing a 
slight curvature in its distal part. The exserted part of the spine is ornamented with 
enamel ridges. One ridge forms a keel on the anterior border of the spine. On the 
proximal part of the spine there are two or three ridges near the anterior border, the 
posterior part of the lateral surface being smooth. Towards the tip, the number of 
ridges increases to about five, equally spaced on the lateral surface. The ridges are 
straight and do not anastomose or bifurcate. The denticles on the posterior face of 
the spine are in a single series, although towards the base they may be placed alter- 
nately to the right and left of the mid-line, indicating their origin from paired series. 

GEOL. II, 7 30 



33° 



BRITISH WEALDEN SHARKS 



I have seen no examples of the double or rudimentary paired denticles of the type 
described in H. hrevicostatus (p. 308). The denticles are smooth and without striae : 
at the tip of the spine they are low and long based, proximally the bases are shorter 
and the denticles higher and recurved. In no specimen is a complete series of 
denticles preserved, but there cannot have been more than about twenty or twenty- 
five. In histological structure (PI. 3, fig. 4) the spines consist of the usual outer layer 
of osteodentine with an inner layer of lamellar tissue in the exserted part of the spine. 
The lamellar tissue makes up from half to two-thirds of the wall of the spine at about 
the middle of its length. 

These spines agree with those which Estes (1964 : 11) assigns to the late Cretaceous 
L. selachos, particularly in the lack of curvature and the form and position of the 
posterior denticles, but they differ in retaining a few enamel ridges on the lateral 
surfaces. 



The Affinities of Lonchidion 

The only other species of Lonchidion known is L. selachos Estes (1964) from fresh- 
water deposits of late Cretaceous age in Wyoming, where isolated teeth are fairly 
common at a niunber of localities. The lateral teeth of L. selachos (Text-fig. 27A) 
are very like those of L. breve breve in the smooth crown, large hybodontoid root. 




&:r^ 



r\ 



i'. 




Fig. 27. Lonchidion selachos Estes. a. Lateral tooth (the holotype) in labial (i), occlusal (2) 
and lingual (3) view, XQ. b. Symphysial tooth in lingual (i) and occlusal (2) view, xii. 
c. Symphysial tooth in basal view, xii. d. Parasymphysial tooth in lingual view, xii. 

E. Cephalic spine, the base restored, in lateral view, x 3-5, and (inset) in dorsal view, x f ■ 

F. Fragment of dorsal fin spine in lateral (i) and posterior (2) view, x 2. All from the Lance 
Formation ; Wyoming, U.S.A. After Estes (1964). 



BRITISH WEALDEN SHARKS 331 

waisted root/crown junction, labial process, and histological structure, but they have 
the occlusal crest produced into fairly well marked cusps ; teeth of L. breve breve 
tending towards this type occur high in the Wealden (see p. 316). In size they 
range from 2-6 mm., a little larger than L. breve. Estes suggests that this type of 
tooth occupied only the lateral and posterior parts of the mouth, and that the anterior 
teeth were of the type shown in Text-fig. 27B, c, d, high-crowned teeth with one or two 
pairs of lateral cusps, no labial process, and squatinoid roots (Casier 1947a : 10) with 
a central canal. In the British Wealden and Purbeck, although several hundred 
teeth of Lonchidion have been examined, no teeth of this type have been found. 
Although it is possible that in Lonchidion the anterior teeth could, by the Upper 
Cretaceous, have evolved a squatinoid root, in parallel with the heterodontids, it 
seems equally probable that the teeth described by Estes belong to a squatinid or 
orectolobid, perhaps to his species Squatirhina americana or a related form. 

Estes compares the lateral teeth of L. selachos with teeth from the German Keuper 
described as Palaeobates spinosus by Seilacher (1943, text-figs. 27, 28). The lateral 
teeth of P. spinosus are typical of the genus, flattened Heierodontus-like teeth which 
bear little resemblance to Lonchidion except in their histological structure. But 
Seilacher assigns to the posterior part of the jaws of P. spinosus very small teeth 
(less than i mm. in length) which are very like those of Lonchidion, with a strong 
labial process, a waisted root/crown junction and a smooth crown. As Estes notes, 
no teeth of this type occurred in Stensio's (1921 : 35) material of associated Palaeobates 
teeth from the Trias of Spitsbergen, and the systematic position of these small teeth 
is by no means certain. Teeth of this type are clearly fairly abundant in the German 
Trias : acid treatment of a small block of bone-bed from the Muschelkalk of Crail- 
sheim (28466) has yielded several isolated crowns which are almost indistinguishable 
from Lonchidion breve breve. Until more complete information on these Triassic 
forms is available it is impossible to decide whether they are related to Lonchidion 
or are convergent, particularly since no Lower Jurassic teeth resembUng Lonchidion are 
known. Estes also draws attention to the resemblance between the teeth of Low- 
chidion and those of the only other freshwater hybodont, the Triassic Lissodus 
africanus (Broom) (Brough 1935, pi. 2). 

Ghkman (1964) has recently proposed a radical reclassification of selachians. 
He divides the sharks and rays among two infraclasses, Orthodonta and 
Osteodonta, mainly on the basis of the histological structure of the teeth. 
In Orthondota the crown of the tooth consists of orthodentine, in Osteodonta of 
osteodentine surrounded by pallial dentine. Glikman finds these two groups to 
have been distinct since their first appearance in the Devonian. He places the 
hybodontids and ptychodonts in the Osteodonta, and makes a new family Polyacro- 
dontidae, order Polyacrodontida and superorder Polyacrodonti within the Orthodonta 
for Polyacrodus and Palaeobates. Thus Glikman believes that Polyacrodus and 
Palaeobates on the one hand and the hybodonts on the other represent lines 
which have evolved independently since the Devonian and are only very distantly 
related. Because of the structure of the teeth, both Lonchidion and Lissodus would 
be placed in the Orthodonta, presumably in the Polvacrodonti, though the regular 



332 BRITISH WEALDEN SHARKS 

alternation of the teeth in adjacent series which occurs in Lissodus and in some 
species of Lonchidion (see p. 319) is not a character of Polyacrodonti according to 
GHkman. Thus Ghkman's major subdivision of the sharks makes it necessary to 
beheve that Lissodus and Lonchidion are only related to the well known Jurassic and 
Lower Cretaceous species of Hybodus in so far as the Orthodonta and Osteodonta 
shared a hypothetical common ancestry in the Lower Devonian. Yet Lissodus and 
Hybodus share such characters as fin spines of exactly similar form and structure, the 
anterior spine lying more obliquely than the posterior (Brough 1935), a posteriorly 
placed anal fin, the presence above the eyes, in males only, of two pairs of cephalic 
spines of similar form, etc. That such features as these should have been acquired 
independently and roughly simultaneously in two unrelated groups seems extremely 
unhkely. Another selachian group in which teeth composed either of osteodentine 
or orthodentine occur in closely related forms is the Upper Cretaceous sub-family 
Ganopristinae of the pristid sawfishes (Schaeffer 1963). In the ganopristines the 
rostral teeth of forms like Onchopristis , Sclerorhynchus and Ganopristis have a crown of 
orthodentine, while Onchosaurus and Pucapristis (like the Tertiary and living Pristi- 
nae) have a crown of osteodentine. Close relationship between these various genera 
seems beyond doubt. I am therefore unable to accept Glikman's Orthodonta and 
Osteodonta, at least so far as the Mesozoic sharks are concerned. I believe that the 
thickness of the pallial dentine and the presence or absence of osteodentine are features 
which have changed a good deal in the history of the sharks, that these characters 
cannot be used to define major subdivisions, and that there are as yet no satisfactory 
grounds on which forms such as Polyacrodus, Palaeobates, Lissodus and Lonchidion 
can be separated from the Hybodontidae. At present, all that can be said of the 
origins of Lonchidion is that the fin spines and cephaUc spines ally it with the Hybo- 
dontidae, the histological structure of the teeth is like Polyacrodus, and that the teeth 
in the earliest known species, L. heterodon, are more like those of Hybodus than they 
are in later species. Similar forms (Lissodus) were already present in fresh water in 
the Trias. 

Lonchidion is evidently a genus of hybodont which became adapted to life in 
fresh water in or before the Upper Jurassic, and underwent considerable radiation 
there, producing Polyacrodus-like forms (L. striatum) and highly specialized forms 
in which the roots of the teeth were lost {L. rhizion), both these being short-lived, 
while the more generalized types [L. breve, L. selachos) continued through almost to 
the end of the Cretaceous as the last survivors of the hybodonts. The relationships 
between the species of Lonchidion are summarized in Text-fig. 31. 

Family PTYCHODONTIDAE 

Amended diagnosis. Hybodont sharks in which there are no fin spines or 
cephalic spines ; vertebral centra possibly calcified ; jaws elongated, with teeth 
confined to their broad symphysial region ; teeth flattened and crushing, dentition 
heterodont ; nine or less paired files of teeth and an unpaired symphysial file in each 
jaw ; largest and most specialized teeth, which are rhombic or rectangular, at or 



BRITISH WEALDEN SHARKS 333 

near the symphysis, teeth decreasing in size posteriorly ; crowns of teeth with strong 
ridges or crests of enamel ; crown made up mainly of osteodentine ; roots of teeth 
hybodontoid. 

Genus HYLAEOBATIS Smith Woodward 1916 : 19 

Amended diagnosis. Ptychodont sharks known only by isolated teeth ; jaws 
probably very broad, dentition probably similar in both jaws ; a symphysial file and 
eight paired files of teeth in each jaw ; symphysial teeth rectangular, three or four 
times as long as broad, parasymphysial teeth rectangular, about two-and-a-half 
times as long as broad ; second paired lateral teeth rhomboid, remaining teeth more 
or less ovoid and Acrodtis-like, not so closely apposed as anterior teeth ; crowns of 
teeth with occlusal crest near labial margin in anterior teeth, central in posterior 
teeth ; fine bifurcating and anastomosing striae radiating from occlusal crest ; 
histological structure of crown like that in Acrodtis, vascular canals less regularly 
arranged than in Ptychodus. 

Type species. Hylaeobatis ornata (Smith Woodward), the only species. 

Hylaeobatis ornata (Smith Woodward) 
(PI. 4 ; PI. 5, figs. 4-7 ; Text-figs. 28-30) 

1889 Acrodus ornatus Smith Woodward : 296, pi. 13, fig. 10. 

1916 Acrodus ornahis Smith Woodward ; Smith Woodward : 14, pi. 2, figs. 15-18. 

1916 Hylaeobatis problemaiica Smith Woodward : 19, pi. 5, figs. 1-5, text-fig. 10. 

Diagnosis. As for genus ; symphysial teeth reaching about 15 mm. in length. 

HoLOTYPE. P. 5275, posterior tooth without root, probably from the Wealden 
Shales, Brixton, Isle of Wight. 

Material. In addition to the holotype, about 300 teeth, all but ten without roots. 

Horizons and localities. Weald Clay : Henfield, Sussex ; Crowhurst, Surrey ; 
Bexhill, Sussex ; Meadvale, Redhill, Surrey ; Hastings, Sussex ; Sevenoaks, Kent. 
Wealden Shales : Brook, Brixton, Yaverland and Atherfield Point, Isle of Wight. 
Lower Greensand : Perna Bed, Sandown, Isle of Wight ; Sandgate Beds, Godalming, 
Surrey (derived). 

Description. The new material of this species from Henfield, consisting of almost 
three hundred teeth (mostly without roots), allows one to attempt a restoration of 
the dentition and shows that the two species Acrodus ornatus and Hylaeobatis 
problematica are synonymous, the first having been established for the small posterior 
teeth and the second for the large anterior teeth. 

The material from Henfield includes nine complete teeth in which the roots are 
preserved : the dimensions of these teeth are shown in Table II. Among these nine 
complete teeth, seven distinct types are present, the two pairs P. 47212, P.47218 and 
P.47213, P.47217 being of the same type. Two more types of tooth are represented 
among the incomplete teeth. 



334 BRITISH WEALDEN SHARKS 

s 




Fig. 28. Hylaeobatis ornata (Smith Woodward). Restoration of the dentition oi one jaw in 
occlusal view, x 1-5 approx. ' s ', S3Tnphysial file ; I-VIII, paired files. The surface pattern 
of one tooth in the symphysial file and each of the right hand files is drawn from examples 
from the Weald Clay of Henfield, Sussex. 

1. Symphysial teeth. The largest and broadest of the complete teeth, P.47211, 
(PI. 4, fig. I ; Text-fig. 28 ' S ') is perfectly bilaterally symmetrical and has a clearly 
marked ridge in the centre of the basal surface of the root : this must be a symphysial 
tooth, and the ridge marks the sjnnphysis between the two rami of the jaw. In 
shape the tooth agrees with a worn specimen figured by Smith Woodward (1916, 
pi. 5, fig. 3), and among the teeth from Henfield there are nine rootless examples of 
this type, ranging in length from 7-15 mm., and in the ratio of breadth to length 
from 2-8-3-5. The labial face of the crown is strongly convex in the vertical plane, 
weakly concave in the horizontal. The lingual face is strongly concave vertically 
and weakly convex horizontally. The lateral ends of the crown are rounded in 
P.47211, but in other specimens they may show two pressure scars, giving a hexagonal 
coronal surface. In P.47211 the crown is quite unworn. The coronal surface is 
divided into a narrow labial zone and a broad lingual zone by a clearly marked 
longitudinal ridge. Comparison with the less specialized posterior teeth shows that 
this ridge is the occlusal crest. The labial zone, which makes up about a quarter of 
the surface of the crown, slopes sharply downwards, and is ornamented with bifurcat- 
ing ridges which pass out from the occlusal crest to end about half way between 
this crest and the root/crown junction. The lower half of this sloping labial surface 
of the crown fitted in life in the hollow on the lingual face of the crown of the preceding 
tooth. The broad lingual zone of the crown is almost flat, and formed the occlusal 
surface of the tooth. It is ornamented with striae which originate at the occlusal 
crest and which bifurcate and anastomose frequently as they pass lingually, so that 
the lingual half of the crown has a reticular surface. The strongly concave lingual 
face of the crown is separated from the coronal surface by a sharp angle, and is 
smooth except for coarse irregularities near the root /crown junction. 

2. Parasymphysial teeth. P.47212 (PI. 4, fig. 2 ; Text-fig. 28, I) is a tooth in 
which the crown is elongated, broad and flat, as in the symphysial teeth, but is 
clearly asymmetrical and is shorter and proportionally broader than in the symphysial 
teeth. As one would expect, teeth of this type are about twice as common as the 
symphysial teeth : there are about twenty incomplete examples among the material 
from Henfield. The tooth figured by Smith Woodward (1916) in PI. 5, fig. 4, is a worn 



BRITISH WEALDEN SHARKS 



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336 BRITISH WEALDEN SHARKS 

example of this type. These teeth range from 6-12 mm. in length, and in the ratio of 
breadth to length from 2-6 to 3-0. The crown is shaped like that of the symphysial 
teeth, being weakly concave labially and convex lingually in the horizontal plane, with 
the occlusal crest near the labial margin, a steeply sloping labial zone, and similar 
ridged and reticulate ornament. The crown is deeper at one end than at the other, 
and the deeper end of the crown commonly bears a pair of pressure scars caused by 
contact with the teeth in the adjacent file. The lower end of the crown is often 
without pressure scars, as in the specimen figured by Smith Woodward. These 
teeth are almost certainly parasymphysial teeth from the first paired files in the 
jaws, the deeper end of the crown being medial and the pressure scars at this end 
being caused by contact with the large symphysial teeth. 

3. Antero-lateral teeth. P.47213 (PL 4, fig. 3 ; Text-fig. 28, II) is an example of 
a third and very distinctive type of tooth, in which the crown is short and very broad, 
with a ratio of breadth to length often less than 1-5, and the total depth almost equal 
to the length of the crown. The crown of this type of tooth is strongly asymmetrical. 
The material from Henfield contains about twenty-five teeth of this type, ranging in 
length from 4-8 mm. and in the ratio of breadth to length from i-3-i-8. The holo- 
type of H. problematica (Smith Woodward, 1916, pi. 5, fig. i) and two more of Smith 
Woodward's figured specimens of this species (1916, pi. 5, figs. 2, 4) are of this type. 
In these teeth the ornamentation of the crown is very like that in the symphysial 
and parasymphysial teeth, but the occlusal crest does not lie so near to the labial 
margin of the crown, and there is often an area of reticulate ornamentation lying 
labial to this ridge, a feature which is not seen in symphysial and parasymphysial 
teeth. As in the parasymphysial teeth, one end of the crown is deeper than the 
other : this deeper end is commonly marked by pressure scars whereas the shallow 
end is almost always without pressure scars. The concavity on the lingual face of 
the crown, in which the labial margin of the succeeding tooth fitted, is much stronger 
at the deeper end of the crown than at the shallower. These facts show that the 
deeper end of the crown is medial, that these teeth were in close contact medially 
with their neighbours in both the same and the adjacent file, but that laterally these 
contacts, particularly with teeth in the adjacent file, were less close. The teeth are 
probably members of the second paired files in the jaws, next to the parasymphysials, 
the two types of teeth being equally common. These antero-lateral teeth show the 
strongest asymmetry of all the teeth. 

4. Lateral and posterior teeth. The three types of teeth described so far, the 
symphysial, parasymphysial and antero-lateral, all have the crown approximately 
rectangular or rhomboid in shape, with a broad, flat coronal surface and with the 
occlusal crest near the labial margin. The remainder of the teeth from Henfield 
do not show these features : the crown is ovoid in shape, its surface is more or less 
rounded, and the occlusal crest lies at or near the centre of the crown. The distinc- 
tion between these two classes of teeth is not absolute, intermediates between the 
two being found in some of the antero-lateral teeth, where the crown may be moder- 
ately rounded and have the occlusal crest some distance from the labial margin, and 
in some of the larger lateral teeth. These intermediate forms and the relative 



BRITISH WEALDEN SHARKS 337 

abundance of the different types of teeth leave Uttle doubt that the two classes of 
teeth are correctly placed in the same species. The symphysial, parasymphysial 
and antero-lateral teeth, interpreted as representing five files (one median and two 
paired) make up a little less than one-third of the sample from Henfield (68 out of 
215 teeth) : this proportion gives some support to the conclusion, reached on morpho- 
logical grounds, that there was a total of seventeen files of teeth in each jaw. 

P. 47214 (PL 4, fig. 4 ; Text-fig. 28, IV) is an example of one of the larger lateral 
teeth. The crown is approximately ovoid in outline. The occlusal crest of the 
crown, though not strongly marked in this specimen, lies nearer to the centre of 
the crown than to the labial margin. The ornamentation of the crown consists of 
bifurcating and anastomosing striae, diverging from the occlusal crest and forming a 
reticulum towards the margins of the crown, including a reticular area towards the 
labial margin, as in the antero-lateral teeth. The material from Henfield contains 
about fifty teeth agreeing with this specimen, ranging in length from 4-0-7-5 mm. and 
in the ratio of breadth to length from i •9-2-7. Like the parasymphysial and antero- 
lateral teeth, these specimens are asymmetrical, with the medial end of the crown a 
little higher than the lateral. Pressure scars on the ends of the teeth are rare, and 
these teeth were evidently not closely associated with the neighbouring files. These 
teeth are interpreted as forming the third and fourth paired files of teeth in each jaw. 

Among the remaining teeth from Henfield, two more distinct types can be easily 
recognized, a long, narrow type, and a short, broad type, both including only small 
teeth. The long, narrow teeth (P.47216, PI. 5, fig. 6 ; Text-fig. 28, VHI) range in 
length from 3- 0-5-0 mm., with a ratio of breadth to length between 2-7 and 3-2. The 
crown is an elongate ovoid in shape. In the larger examples the ornamentation of 
the crown is irregular, though an occlusal crest in the centre of the crown can normally 
be recognized, from which bifurcating and anastomosing ridges diverge, forming a 
reticular pattern towards both lingual and labial margins of the crown. The occlusal 
crest curves lingually at each end of the crown. In smaller teeth of this type the 
ornamentation is more regular, with a strongly marked occlusal crest which curves 
lingually at the ends of the crown and forms a weak central cusp in the centre. The 
lingual half of the coronal surface is almost smooth in the smallest teeth, but usually 
bears a ridge, running parallel to the occlusal crest, which is joined to the central 
cusp and curves lingually without reaching the ends of the crown. The labial half 
of the crown bears two or three similar longitudinal ridges, joined to the central 
cusp and turning labially before reaching the ends of the crown. The holotype of 
Acrodus ornatus Smith Woodward (P. 5275 ; 1889, pi. 13, fig. 10) is a worn tooth of 
this type, and among the material from Henfield there are about twenty-five 
examples. 

The short, broad teeth (Text-fig. 28, VII) in none of which is the root preserved, 
agree with the long narrow teeth described above in all features of ornamentation, 
but are typically 3-4 mm. in length and 1-6-2-1 mm. in breadth, with a ratio of 
breadth to length of i-6-i-g. They show the same increasing regularity of orna- 
mentation with decreasing size, with the appearance in small forms of ridges running 
parallel to the occlusal crest. The crown in these teeth is almost semicircular in 



338 BRITISH WEALDEN SHARKS 

shape, with a flat hngual margin and a strongly arched labial one. A tooth of this 
type is figured as Acrodus ornatus by Smith Woodward (1916, pi. 2, fig. 18), and 
there are about thirty examples among the material from Henfield. 

There remain undescribed about fifty teeth from Henfield, which are intermediate 
in shape and ornament between the large lateral teeth (III, IV) and the small teeth 
of short, broad (VII) and long, narrow (VIII) type. In these teeth (P.47215 ; 
PL 5, fig. 7) the crown is ovoid in shape, 3-o-7-o mm. in length, and with a ratio of 
breadth to length of i-9-3-o. The larger of these, 4-5-7-0 mm. in length, are narrower, 
with the ratio of breadth to length usually between 2-5 and 3-0, while the smaller 
teeth, 3-0-4-5 mm. in length, are broader, with the ratio of breadth to length from 
2-0 to 2-5. The labial margin of the crown is usually more strongly curved than the 
lingual. An occlusal crest is normally recognizable in the centre of the crown, with 
bifurcating and anastomosing striae diverging to form a reticular pattern at both 
lingual and labial margins. One specimen figured as Acrodus ornatus by Smith 
Woodward (1916, pi. 2, fig. 17) is a worn tooth of this type. Of each of the paired 
types of teeth described above, the material from Henfield normally contains between 
twenty and thirty examples. This material contains about fifty of these ovoid teeth, 
and they probably represent two paired files in the postero-lateral part of the jaw 
(Text-fig. 28, V-VI). 

5. The Root. There is no significant variation in the shape or structure of the 
root between the small posterior teeth and the large sjmiphysial teeth. In all the 
teeth the root is typically hybodontoid (Casier I947fl : 9) in structure : porous and 
trabecular, with an irregular series of large foramina in both the lingual and labial 
surfaces (PI. 4) . The labial face of the root is concave, and is clearly marked off from 
the flat basal surface, which slopes lingually. The lingual face of the root is strongly 
convex, its upper part sloping out in a ledge which is largest in the anterior teeth, 
where it fitted against the concavity on the labial face of the root of the succeeding 
tooth. The root is shorter and narrower than the crown in all but the smallest 
posterior teeth (PI. 5, figs. 6, 7). In the posterior teeth the root is shallower than 
the crown, but in the large anterior teeth it may be as deep as or deeper than the crown. 

6. Histological Structure. Smith Woodward (1916 : 20, text-fig. 10) has given 
a brief account of the histological structure of the crown, which he compares with 
that of Ptychodus. Sections of the crowns of a large anterior tooth and a small 
posterior one are shown in PL 5, figs. 4, 5. The structure of the crown seems to be 
closer to that of Acrodus (Owen 1840, pi. 14), with which it agrees in the rather sparse 
and irregularly arranged vascular canals and the ' bunching ' of the tubules of the 
thin pallial dentine where they arise from the vascular canals. In Ptychodus the 
vascular canals are more regularly and closely arranged, and are parallel throughout 
the distal part of their length, as they are in the teeth of rays (Casier 1953, pis. 1,2). 

7. Other remains. There are no shark vertebrae associated with the abundant 
teeth of Hylaeobatis at Henfield, suggesting that the notochord was uncalcified, as in 
the hybodontids, or that the centra were very weakly calcified. As all the fin spines 
from Henfield are of normal hybodont type and seem to belong either to Hybodus 
basanus or to Lonchidion, there is no evidence that fin spines were present in Hylaeobatis. 



BRITISH WEALDEN SHARKS 339 

Restoration of the dentition. On the basis of the morphology and relative abundance 
of the various types of teeth described above, a tentative reconstruction of the jaw 
of Hylaeobatis ornaia has been made (Text-fig. 28). This reconstruction is largely 
hypothetical in that there is, as yet, no means of distinguishing between teeth from 
the upper and lower jaws, and one cannot discover whether there was any difference 
between the arrangement of the teeth in the two jaws, as there was in Ptychodus. 
But since only one type of symphysial tooth has been found, which when worn has a 
single, centrally placed wear facet, any variation between the two jaws is unlikely 
to be of the type seen in Ptychodus, where the sjonphysial teeth are large in one jaw 
and small in the other (Smith Woodward 1888 : 296). Smith Woodward (1916 : 21, 
pi. 5, fig. 4) has described one antero-lateral tooth of Hylaeobatis as showing signs of 
having been opposed by two teeth in life, but this is not true of any of the teeth from 
Henfield, and at present there is no good evidence of dissimilarity between the two j aws. 

Since the symphysial teeth are the largest teeth, with the flattest crowns, the most 
labially displaced occlusal crest, and the closest fit with their neighbours in the same 
and adjacent files, the very broad, weakly curved dentition suggested in Text-fig. 28 
seems the most reasonable interpretation of the material. As the crowns of the 
symphysial and parasymphysial teeth are slightly concave forwards, the medial 
part of the dentition must have been transversely placed and without appreciable 
posterior curvature. The strongly asymmetrical antero-lateral teeth (II) must mark 
the transition between the transversely placed, pavement-like dentition of the front 
of the mouth and the more Acrodus-like teeth of the last six files, which were probably 
curved back towards the articular region of the jaws. The asymmetry of the 
antero-lateral teeth and the close fit with the parasymphysial teeth which is indicated 
by the pressure scars on their medial faces show that these teeth must have curved 
forwards, as shown in Text-fig. 28. Because of this, it is difficult to see how the 
curvature of the posterior part of the jaw could have been greater than that shown 
in the figure, if the files of teeth were in reasonably close contact. All this suggests 
that the jaws of Hylaeobatis were very broad anteriorly, and that the dentition was 
largely confined to the anterior, transverse part of the jaw, as more complete remains 
show that it was in Ptychodus (Smith Woodward 1904). The transition between 
the small, AcrodiisA\\it posterior teeth and the large, flattened teeth at the symphysis 
shows how a ray-like dentition can be produced in a hybodont by specializations which 
include the shifting labially of the occlusal crest, so that the occlusal surface is formed 
by the lingual face of the crown, the hollowing out of the lingual face of the crown to 
give increased contact between successive teeth, and the development of a rough 
surface by anastomosis between the surface striae. The variability in the occurrence 
and size of the pressure scars on the lateral and medial surfaces of the crowns of the 
anterior teeth suggests that in Hylaeobatis, just as in Ptychodus, the teeth in adjacent 
files did not alternate regularly in position as they do in the rays. 

The Affinities of Hylaeobatis 

Smith Woodward (1916 : 19) placed Hylaeobatis in the Myliobatidae, but compared 
it with Ptychodus (which he then included in the same family) and suggested that it 



340 BRITISH WEALDEN SHARKS 

might be intermediate between Ptychodus and the cestracionts (including hybodonts) . 
Later (1932 : 83) he included the genus in the Ptychodontidae. Casier (1953) has 
discussed the origin and affinities of the ptychodonts in some detail, and has shown 
that they are almost certainly specialized derivatives of the hybodonts or hetero- 
dontids, probably the former. He considered (p. 34) Hylaeobatis to be as close to 
the heterodontids as to the ptychodonts, and saw in it ' une forme intermediaire, au 
point de vue de la morphologie dentaire, entre les Hybodontiformes et les Ptycho- 
dontes '. More recently (1961 : 45, pi. 5, fig. i), in describing as Hylaeobatis? sp. a 
tooth from the Neocomian of the Congo, Casier revised this opinion and concluded 
that Hylaeobatis is probably a pycnodont. The tooth described by Casier seems to 
bear little resemblance to Hylaeobatis, particularly in the ornamentation of the occlu- 
sal surface and the flat, ornamented lingual face, and is probably a pycnodont tooth. 

The new material described here seems entirely to confirm Smith Woodward's 
and Casier's (1953) opinions on the affinities of Hylaeobatis. Hylaeobatis shows 
unmistakable signs of hybodont relationships in the Acrodus-like posterior teeth 
and histological structure of the crown, and in the hybodontoid roots of the teeth. 
On the other hand, there are equally important indications of relationship with 
Ptychodus. Casier (1953 : 25) considers that the ptychodont type of dentition evolved 
from hybodonts with undifferentiated dentition by specialization of the anterior 
teeth and reduction in the number of files of teeth (in contrast to the heterodontids, 
where the lateral teeth are specialized, the anterior teeth remaining small) . Hylaeo- 
batis exhibits the most important character of the ptychodonts, the enlargement and 
specialization of the anterior teeth, with the symphysials and parasymphysials the 
most specialized. As to the number of files of teeth, Ptychodus decurrens, which 
seems to be the least specialized of the well known species (see below), has only five 
or six paired files in addition to the symphysials, but in later species the number of 
paired files rises (perhaps secondarily) to eight in P. mortoni (Williston 1900 : 238) 
and to nine in the Upper Senonian P. mediterraneus (Canavari 1916 : 37). Hylaeo- 
batis, with eight paired files and a symphysial, agrees with P. mortoni and with 
hybodontids such as Acrodus curtus {=A. anningiae) (Smith Woodward 1889, 
text-fiig. 10). The number of paired files is rather variable in hybodontids (9 or 10 
in H. basanus, 9 in H. brevicostatus , six in Aster acanthus), and too much importance 
should not be attached to it. In the apparent absence of fin spines Hylaeobatis 
agrees with Ptychodus and differs from the hybodontids, in the apparent absence of 
calcified centra it agrees with the hybodontids (the evidence for the presence of 
calcified centra in Ptychodus is not conclusive — see below). 

There seems little doubt that Hylaeobatis is intermediate between the hybodontids 
and Ptychodus. The genus is placed in the Ptychodontidae because it already 
exhibits the specialization of the anterior teeth which is characteristic of this family. 
There are two further points to be considered : the origin of Hylaeobatis and the 
ptychodontids, and the nature of the relationship between Hylaeobatis and Ptychodus. 

The Origin of Hylaeobatis 

Casier argues that forms with a ptychodont dentition, with the anterior teeth the 
largest and most specialized, must have evolved from hybodontoids in which the 



BRITISH WEALDEN SHARKS 



341 



dentition was homodont, without the enlargement of the lateral teeth which is 
characteristic of most hybodontids and of heterodontids. Also, since the anterior 
teeth of Hylaeohatis are the most specialized, it seems reasonable to assume that the 
small posterior teeth will retain most resemblance to the ancestral form. In the 
Weald Clay at Henfield, where teeth of Hylaeohatis are very abundant, there is an 
almost perfect gradation between the smaller posterior teeth of this genus and teeth 
of Lonchidion breve breve, a hybodontid species in which the dentition was probably 
almost homodont (see p. 314). In the smallest teeth of Hylaeohatis (Text-fig. 29c), 
possibly juvenile, the surface ornamentation is reduced to a single central occlusal 
crest with a few weakly marked cusps along it, and a single labial accessory cusp 
overlying a broad labial process. Such teeth are similar in shape to but broader than 
teeth oi Lonchidion breve : they are particularly like L. breve pustidatum (Text-fig. 19), 
a subspecies known only from the Perna Bed at the base of the Atherfield Clay. 
The histological structure of the teeth of Lonchidion breve, with their very thick 







2 mm 

Fig. 29. A-c. Posterior teeth of Hylaeohatis ornata (Smith Woodward) from the Weald Clay 
of Henfield, Sussex, in occlusal view, to show the transition from normal posterior teeth (a) 
to (?) juvenile teeth (c) which resemble Lonchidion. P. 47268-70. d. Transverse section of a 
small posterior tooth crown of Hylaeohatis ornata, P.47278, Weald Clay ; Henfield, Sussex. 
E. Transverse section of a tooth crown of Lonchidion breve breve sp. nov., P.47279, Ashdown 
Beds, Cliff End Bone-bed ; Cliff End, Sussex. 



342 BRITISH WEALDEN SHARKS 

pallial dentine, seems at first to be against any relationship with Hylaeohatis, in 
which the crown consists mainly of osteodentine with a rather thin layer of pallial 
dentine whose tubules arise in bunches from the terminal parts of the vascular 
canals of the osteodentine (see p. 331 for discussion of Glikman's (1964) views on the 
differences between these types of tooth structure). But the differences here seem 
to be due largely to the greater size and thickness of the crown in Hylaeohatis. In 
Lonchidion the tubules of the pallial dentine arise from the tips of vascular canals, 
just as in Hylaeohatis, but the crown is so low and narrow that there is normally 
only a single row of vascular canals at the base of the crown : other vascular canals 
are sometimes present (Text-fig. 29E), from which a few short dentine tubules are 
given off into the basal parts of the crown. In the smallest teeth of Hylaeohatis 
(Text-fig. 29D) the structure may be very like that of Lonchidion : in these teeth 
the vascular canals only enter the base of the crown, there giving off sprays of 
pallial dentine tubules which make up the bulk of the crown. The type of structure 
seen in the teeth of Hylaeohatis can be derived from that of Lonchidion by increase 
in the breadth of the crown, making more vascular canals and tufts of palHal tubules 
necessary, and increase in the depth of the crown without increase in the thickness of 
the pallial dentine, allowing the basal part of the crown to be formed by osteodentine 
laid down around the vascular canals. 

At present, I would suggest that Hylaeohatis (and the Ptychodontidae) arose from 
a homodont species of the hybodontid genus Lonchidion, primarily by reduction in 
the labial process, by increase in the surface ornament, by specialization in the 
anterior teeth, and by loss of fin and cephalic spines. 

Relationships within the Ptychodontidae 

The Ptychodontidae contains only the three genera Ptychodus, Hylaeohatis and 
Heteroptychodus. Heteroptychodus Yabe & Obata (1930 : 6), type species H. stein- 
manni Yabe & Obata (1930 : 7, pi. 2, figs. 6-8), is known only by a single tooth crown 
from the basal Cretaceous of Japan. This tooth is quite unhke Hylaeohatis and 
rather unlike Ptychodus, but the genus is so poorly known that nothing is to be 
gained from discussing it. I agree with Smith Woodward (1912 : 245) that the 
separate genus Hemiptychodus Jaekel (1894 : 137) for P. mortoni Mantell, in which 
the striae radiate from the centre of the tooth crown, is not justified. 

Since Hylaeohatis is unknown above the base of the Aptian and Ptychodus does 
not appear until the Cenomanian there is a long gap in the history of the Ptycho- 
dontidae, and the two genera are not necessarily closely related. There is a number 
of characters in which Ptychodus is more speciahzed than Hylaeohatis, including : 

I. Differences between the dentition of the two jaws in Ptychodus, one having 
very small symphysial teeth and large parasymphysials, the other having the largest 
teeth on the symphysis. Smith Woodward (1904), on the basis of a specimen in 
the Willett Collection in Brighton Museum in which the greater part of the cartilage 
of one jaw and fragments of the other are preserved, interpreted the jaw with the 
small symphysial teeth as the upper, but Canavari (1916 : 92) has shown that these 
small symphysial teeth were sunk beneath the level of the parasymphysials so that 



BRITISH WEALDEN SHARKS 343 

they were not functional but formed the floor of a median groove in the dentition. 
He argues that such a groove v^^ould be without function in the upper jaw, but that 
in the lower it would serve as a gutter down which masticated food would be washed 
to the oesophagus. This interpretation seems reasonable and may provide an 
explanation for this difference between Ptychodus and Hylaeobatis , in which there 
is no evidence of differentiation between the upper and lower dentition. 

2. Differences in the histological structure of the teeth. In Ptychodus (Owen 
1840, pis. 18, 19 ; Casier 1953, pi. i) the vascular canals of the crown are long, 
straight, and parallel through the greater part of their length, producing a tissue 
which resembles the tubular dentine of holocephalans and dipnoans. This type of 
tissue is an adaptation to a durophagous diet (Radinsky 1961 : 79) which has arisen 
independently in a number of groups, including rays, Ptychodus and Asteracanthus 
among selachians. The type of structure seen in Ptychodus can probably be derived 
from that in Hylaeobatis by alignment of the ascending vascular canals which give 
off the paUial dentine, in just the same way as the teeth of the hybodontid Astera- 
canthus {Ptychodus-like in structure) are probably derived from those of Acrodus 
{Hylaeobatis-like in structure). 

3. Ptychodus is thought to have had calcified vertebral centra, while there is no 
evidence for such structures in Hylaeobatis. Direct evidence for the presence of 
calcified centra in Ptychodus rests on two specimens, one from the English Chalk 
(39436, P. decurrens ; Smith Woodward 1912, pi. 52, figs. 1-6) in which two centra 
are preserved in association with fragments of calcified cartilage and twenty-three 
scattered teeth, and one from the Italian Upper Senonian, the holotype of P. 
mediterraneus Canavari (19 16), in which an impression of a single centrum is preserved 
in association with an almost complete dentition. In my opinion there is still 
considerable doubt as to the weight to be attached to these specimens. In favour 
of the centra belonging to the same animal as the teeth is the fact that both the centra 
and the teeth in the two specimens are of approximately the same size (the centra 
c. 50 mm. in diameter in both, parasymphysial teeth from the lower jaw c. 35 x 29 
mm. in Canavari's specimen, 34 x 31 mm. in the English specimen) : it is unlikely 
that this should be so if the teeth and centra were associated either by chance or by 
the ingestion of Ptychodus by a shark with calcified centra. Against the centra 
belonging to Ptychodus there are several points. First, as Smith Woodward (1912 : 
229) noted, the centra are very like those found in association with teeth of Squalicorax 
in the Kansas Chalk, where there is no doubt that the teeth and vertebrae are from 
the same animal : I have been unable to find any significant differences between 
the two types of centra in external or internal structure. Secondly, a considerable 
nimiber of associated dentitions of Ptychodus has been collected in the English and 
American Chalk : it seems strange that only in one of these (one of the least complete) 
are centra preserved. Thirdly, calcified centra are not known in any other hybo- 
dontiform shark. It appears that there is not enough evidence to regard the presence 
of centra as a genuine difference between Hylaeobatis and Ptychodus. 

The earliest well-known species of Ptychodus is P. decurrens Agassiz, the only 
species known in the Cenomanian zones of the English Chalk (Dibley 191 1 : 273). 



344 



BRITISH WEALDEN SHARKS 



This species is more like Hylaeobatis than are later species in that there is no broad, 
flat, marginal zone on the crown, and it is therefore probably primitive. As the 
symphysial teeth of Piychodtis are the most specialized, one may expect that, as in 
Hylaeobatis , the small posterior teeth will show most resemblance to the ancestral 
form. A posterior tooth of P. decurrens is shown in Text-fig. 30A. Although larger 
and more strongly ornamented than posterior teeth of Hylaeobatis, it shows the same 
relative proportions of crown and root, and has an approximately similar surface 






5 mm 





1 mm 



1 mm 

Fig. 30. A. Ptychodus decurrens Agassiz. Posterior tooth in labial (above), occlusal (centre) 
and lingual view. 4361, Lower Chalk ; Lewes, Sussex, b. Ptychodus decurrens var. oweni 
Dixon. Parasymphysial tooth, probably from the first paired file of the upper jaw, right 
side, in occlusal view (symphysis to the right, labial margin uppermost). 39125, Lower 
Chalk ; Hailing, Kent. c. Hylaeobatis ornata (Smith Woodward). Antero-lateral tooth 
from the second paired file in occlusal view (symphysis to the left, labial margin uppermost). 
P. 47230, Weald Clay ; Henfield, Sussex. 



BRITISH WEALDEN SHARKS 345 

ornamentation, a longitudinal occlusal crest from which bifurcating striae diverge. 
The arrangement of the striae tends towards the parallel ridges characteristic of 
anterior teeth in Ptychodus, but parallel transverse ridges also tend to develop in 
posterior teeth of Hylaeobatis (p. 337). Differences between teeth of P. deciirrens 
and Hylaeobatis include : 

1. The striae on the crown in Ptychodus extend almost to the root /crown junction 
on all surfaces of the tooth : this seems to be true of all except the smallest posterior 
teeth (Text-fig. 30A). In Hylaeobatis the lingual face of the crown is always smooth 
or nearly so, and on the other faces the striae end well above the root/crown junction. 

2. In Hylaeobatis the whole of the lingual face of the crown is occupied by a 
concavity in which the labial surface of the succeeding tooth fits : this is true even 
of the smallest teeth. In the small posterior teeth of Ptychodus there may be no 
cavity at all, as in Text-fig. 30A, while in the anterior teeth the cavity is always rather 
small and often appears to be more a pressure scar caused by the succeeding tooth 
rather than a real feature of the tooth. In Ptychodus the surface of the lingual 
cavity is always ornamented, like the rest of the crown, while in Hylaeobatis it is 
smooth. This seems to imply that in Ptychodus the fit between successive teeth 
was not so close as it was in Hylaeobatis . 

3. In Ptychodus the crowns of the anterior teeth are almost square or only 
slightly longer than broad. There is no counterpart in Ptychodus of the transversely 
elongated symphysial and parasymphysial teeth of Hylaeobatis, nor is there any sign 
in Ptychodus of the labial shifting of the main occlusal crest which is characteristic 
of these anterior teeth in Hylaeobatis. But the antero-lateral teeth of Hylaeobatis 
(II, Text-fig. 28) are almost identical in shape with parasymphysial teeth of P. 
decurrens, and in the ornamentation of the crown they are very close to P. decurrens 
var. oweni (Smith Woodward 1912, pi. 52, figs. 9-11) a form in which the striae are 
more irregular than they are in typical exampfcs of the species. The teeth shown 
in Text-fig. 30B, c demonstrate this very close smiilarity, the only difference between 
the two being the finer and more extensive orrSmentation of P. decurrens, a feature 
which could well be due simply to the much larger size of the latter. 

In summary, there are some characters in which Ptychodus is more advanced than 
Hylaeobatis, particularly the histological structure of the teeth and the different form 
of the symphysial teeth in the upper and lower jaws, but both these characters could 
have evolved from the conditions in Hylaeobatis. In Ptychodus the crown is more 
heavily and extensively ornamented than in Hylaeobatis, and there was a less close 
fit between successive teeth — in the first of these characters Ptychodus is probably 
more advanced than Hylaeobatis, in the second more generalized, but again these 
conditions could be derived from those in Hylaeobatis. In Ptychodus there is no 
counterpart of the specialized elongated symphysial and parasymphysial teeth of 
Hylaeobatis, but there is a close similarity between the antero-lateral teeth of the 
second paired file in Hylaeobatis and the parasymphysial teeth of P. decurrens. 
This suggests that Ptychodus did not evolve direct from Hylaeobatis, or at least not 
from the only known species of that genus, but from a similar form in which the 
symphysial and parasymphysial teeth were less specialized. 

GEOL. II, 7 31 



346 BRITISH WEALDEN SHARKS 

IV ECOLOGY AND RELATIONSHIPS OF THE FAUNA 

It now seems certain that the bulk of British Wealden (excepting the upper part 
of the Weald Clay and the Wealden Shales : Anderson 1963 ; Casey 1961 : 490) 
and the greater part of the Middle and Upper Purbeck (Anderson 1958) were laid 
down in fresh water (Allen 1959). At Henfield this is confirmed for the fish horizons 
by the ostracods and charophytes (see p. 286). The abundance and variety of the 
shark fauna is therefore surprising, for shark remains are normally taken as evidence 
of marine or estuarine conditions, since among living elasmobranchs only some species 
of Carcharhinus, potamotrygonid rays and pristids have become adapted to life in 
fresh or brackish water. As for the hybodonts, Casier (1961 : y/) is of the opinion 
that they were all marine while Estes (1964 : 167) notes that Lonchidion selachos is 
only the second freshwater form known, the other being Lissodus africanus (Brough 
1935). It is notable that none of the more advanced selachian groups is present in 
the British Wealden and Purbeck, although notidanids, heterodontids, orectolobids 
scyliorhinids, squaUds, squatinids and rhinobatids were already present in Upper 
Jurassic seas : this provides additional evidence that the deposits were laid down in 
fresh water. The moderately large and varied hybodont fauna of the English 
Wealden and Purbeck shows that by the end of the Jurassic (if not before) some 
hybodont sharks had become euryhaline and entered fresh waters. This move 
was clearly advantageous : it removed the hybodonts from competition with more 
advanced forms, some of which (particularly notidanids, heterodontids, orectolobids, 
rhinobatids) must have occupied very similar niches, and allowed them free rein as 
almost the only aquatic predators among the rich teleostean, moUuscan and arthro- 
podan fauna of the Wealden lakes and rivers. The hybodonts were here able to 
undergo a new adaptive radiation, analogous to their marine radiation at their first 
appearance in the Triassic. This radiation, taking place in the absence of other 
selachian competitors, is almost certainly responsible for the similarity between the 
Cretaceous freshwater selachian fauna and the Triassic marine fauna which emerges 
from a list of the species most resembling the Wealden forms. The high-crowned 
Wealden Hybodus species, H. basanus, H. ensis and H. parvidens, resemble similar 
forms which occur throughout the Triassic and Jurassic. The low-crowned H. 
brevicostatus is most like the larger Triassic Polyacrodus species and low-crowned 
Liassic Hybodus such as H. delabechei. Lonchidion breve resembles the Triassic 
Lissodus and Triassic teeth assigned to Palaeobates ; L. striatum resembles the 
smaller Triassic Polyacrodus species ; L. heterodon is another form resembling 
Triassic Palaeobates. Lonchidion rhizion and Hylaeobatis are the only entirely novel 
forms in the Wealden, the first apparently unique, the second leading on to the 
Upper Cretaceous ptychodonts. Other specialized Lower Cretaceous hybodonts 
include the ' hybodontoide de position systematique indeterminee ' described by 
Casier (1961 : 18, pi. 3, figs. 3-6, text-figs. 2, 3) from the Congo, in which there is a 
very deep root with two large canals in the centre and a low, crescentic crown of 
Acrodus-like histological structure. This form, known by isolated teeth from a 
number of localities, is quite possibly another freshwater hybodont. It occurs in 



BRITISH WEALDEN SHARKS 

Hybodus 
i7hngworthi,etc. 




Fig. 31. Diagram showing the probable interrelationships of the sharks of the British Wealden 
and Purbeck and certain Upper Cretaceous species. Lines converging upwards indicate 
convergent evolution, lines converging downwards indicate phylogenetic relationship. Only 
those formations from which sharks are known are included. 

beds of freshwater character (Casier 1961 : 73) which Casier finds to be marine largely 
because Hybodus, which he thought to be exclusively marine, is present (p. yy). 

It seems probable that the hybodonts successful invasion of fresh water in the 
Lower Cretaceous was not permanent : some euryhalinity was retained, and in the 
Upper Wealden some of the more specialized forms were able not only to accommo- 
date to the influx of salt water at the end of the Wealden, but to compete successfully 
with the more advanced selachians present in the sea. The last of the marine hybo- 
dontids (Hybodus illingworthi, Acrodus dolloi, etc., Cenomanian to Senonian) are 
probably derived from the Wealden H. brevicostatus, and the ptychodonts, highly 
successful and widely distributed in the Upper Cretaceous, seem to have evolved from 
near the Wealden Hylaeobatis. 

Within the Wealden and Purbeck, fairly large samples of teeth from successive 
horizons give an unusually complete and well documented account of the evolution 
of the various species. These changes are summarized in Text-fig. 31, the details 
being given in the descriptions of the species. I have no doubt that this diagram is a 
gross simpUfication of the true picture, and that further sampUng will produce 
additions to the fauna and to the complexity of the dendrogram. 



348 BRITISH WEALDEN SHARKS 

This work has been made possible by those who have collected and presented the 
bulk of the new material described : Dr. K. A. Kermack and his colleagues at Univer- 
sity College and Messrs J. F. Wyley, I. M. West, P. J. Whj^brow andB. H. Newi lan. 
I am most grateful to all these gentlemen, in particular to Mr. Wyley for his ca'eful 
collecting at Henfield and for allowing me to accompany him on visits to the pit. 
My thanks are also due to Dr. F. W. Anderson of H.M. Geological Survey, who has 
kindly examined samples of ostracods from Henfield, to Mr. H. A. Toombs for his 
help, and to Mr. P. J. Green, who took the photographs. 

V REFERENCES 

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Agassiz, J. L. R. 1833-44. Recherches sur les Poissons Fossiles. 5 vols. 1420 pp., 396 pis., 
with supplement. Neuchatel. 

Allen, P. 1949. Notes on Wealden Bone-beds. Proc. Geol. Ass., Land., 60 : 275-283, 

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1955- Age of the Wealden in North-Western Europe. Geol. Mag., Lond., 92 : 265-281, 

2 figs. 
1959- The Wealden Environment : Anglo-Paris Basin. Philos. Trans., London (B) 

242 : 283-346, 24 figs. 
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24 : 1-28, 5 figs. 
Anderson, F. W. 1959. Purbeck Beds. In The Geology of the Country around Bridport 

and Yeovil. Mem. Geol. Siirv. U.K. xii + 239 pp., 7 pis. 
1963. Ostracod Faunas in the Weald Clay. In The Geology of the Country around 

Maidstone. Mem. Geol. Surv. U.K., viii -\- 152 pp., 5 pis. 
Berg, L. S. 1955. Classification of fishes and fish-like vertebrates, living and fossil. 2nd edit., 

corrected and enlarged [In Russian]. Trav. Inst. zool. Acad. Sci. URSS, Leningrad, 

20 : 1-286, 263 figs. 
Brough, J. 1935. On the Structure and Relationships of the Hybodont Sharks. Mem. 

Manchr. lit. phil. Soc, 79 : 35-49, pis. 1-3. 
Canavari, M. 1916. Descrizione di un notevole esemplare di Ptychodns Agassiz trovato nel 

calcare bianco della Creta superiore di Gallio nei Sette Comuni (Veneto). Palaeontogr. ital., 

Pisa, 22 : 35-102, pis. 5-14. 
Casey, R. 1961. The Stratigraphical Palaeontology of the Lower Greensand. Palaeontology , 

London, 3 : 487-621, pis. 77-84. 
Casier, E. 1947a. Constitution et livolution de la Racine Dentaire des Euselachii. I. Note 

pr61iminaire. Bull. Mus. Hist. nat. Belg., Bruxelles, 23, 13 : 1-15, 3 figs. 
igi^yb. Constitution et Evolution de la Racine Dentaire des Euselachii. II. fitude 

comparative des types. Bull. Mus. Hist. nat. Belg., Bruxelles, 23, 14 : 1-32, 5 pis. 

1953- Origine des Ptychodontes. Mem. Inst. Sci. nat. Belg., Bruxelles (2) 49 : 1-51, 2 pis. 

1961. Mat6riaux pour la Faune Ichthyologique Eocr6tacique du Congo. Ann. Mus. 

Afr. Cent. 8vo Sci. Geol., Tervuren, 39 : xii + 96 PP-, 12 pis. 
Clemens, W. A. i960. Explanation of an exhibit of specimens of new Wealden mammals. 

Proc. Geol. Soc. Lond., 1588 : 90. 

1963. Wealden mammalian fossils. Palaeontology, London, 6 : 55-69, 10 figs. 

Dalinkevicius, J. A. 1935. On the Fossil Fishes of the Lithuanian Chalk. I. Selachii. 

Mem. Fac. Sci. Univ. Lithuanie, Kaunas, 9 : 245-305, pis. 1-5. 
Dibley, G. E. 191 1. On the Teeth of Ptychodus and their Distribution in the English Chalk. 
/. Geol. Soc. Lond., 67 : 263-277, pis. 17-22. 



BRITISH WEALDEN SHARKS 349 

Egerton, p. M. de G. 1845. Description of the Mouth of a Hybodus found by Mr Boscawen 

Ibbetson in the Isle of Wight. Qtiart. J . Geol. Soc. Land., 1 : 197-199, pi. 4. 
1854. On some new Genera and Species of Fossil Fishes. Ann. Mag. Nat. Hist., London 

(2) 13 : 433-436. 
EsTES, R. 1964. Fossil Vertebrates from the Late Cretaceous Lance Formation, Eastern 

Wyoming. Bull. Dep. Geol. Univ. Calif., Berkeley & Los Angeles, 49 ; 1-187, 5 pis. 
Fritsch, a. 1878. Die Reptilien iind Fische der bohmischen Kreide formation. 46 pp., 10 pis. 

Prag. 
Glikman, L. S. 1964. Akitly paleogena i ikh stvatigraficheskoe znachenie. 229 pp., 31 pis. 

Moskva, Akad. Nauk SSSR. 
Hughes, N. F. 1958. Palaeontological Evidence for the Age of the English Wealden. Geol. 

Mag., Lond., 95 : 41-49, i fig. 
Jaekel, O. 1889. Die Selachier aus dem oberen Muschelkalk Lothringens. Abh. geol. 

Spezialk. Els.-Loth., Strassburg, 3 : 272-332, pis. 7-10. 

1894. Die eocdnen Selachier vom Monte Bolca. \i(y pp., 8 pis. Berlin. 

1898. Ueber Hybodus Ag. S.B. Ges. naturf. Fr. Berl., 1898 : 135-146, 3 figs. 

1906. Neue Rekonstruktionen von Pleuracanthus sessilis und von Polyacrodus {Hybodus) 

haiiffianus. S.B. Ges. naturf. Fr. Berl., 1906 : 155-159, i pi. 
Kermack, K. a.. Lees, P. M. & Mussett, F. 1965 Aegialodon dawsoni, a new tritoberculo- 

sectorial tooth from the Lower Wealden. Proc. Roy. Soc, London (B) 162: 535-554, 

pis. 55-58. 
Leriche, M. 191 1. Sur quelques Poissons du Cr6tac6 du Bassin de Paris. Bull. Soc. geol. Fr., 

Paris (4) 10 : 455-474, pi. 6. 
1929. Les Poissons du Cr6tac6 marin de la Belgique et du Limbourg hollandais. Bull. 

Soc. beige Geol. Pal. Hydr., Bruxelles, 37 : 199-299, 19 figs. 
1930. Rectifications de nomenclature au sujet du grand Cd;rithe du Tuffeau de Ciply 

(Montien) et de ' Hybodus ' de la Glauconie de Lonz6e (Santonien) . Bull. Soc. beige Geol. 

Pal. Hydr., Bruxelles, 39 : 102-105. 
MiLBOURNE, R. A. 1 961. Field Meeting in the Gault at Small Dole, near Henfield, Sussex. 

Proc. Geol. Ass., Land., 72 : 135-138. 
Owen, R. 1840-45. Odontography. Ixxiv + 655 pp., atlas 168 pis. London. 
Peyer, B. 1946. Die schweizerischen Funde von Asteracanthus [Strophodus) . Abh. schweiz. 

paldont., Basel, 64 : i-ioi, pis. i-ii. 
Radinsky, L. 1961. Tooth Histology as a Taxonomic Criterion for Cartilagenous Fishes. 

/. Morph., Philadelphia, 109 : 73-81, pis. i-io. 
Reeves, J. W. 1947. The Henfield Neighbourhood, /w Whitsun Field Meeting to the Central 

Weald. Proc. Geol. Ass., Lond., 58 : 73-85. 

1958. Subdivision of the Weald Clay in Sussex. Proc. Geol. Ass., Lond., 69 : 1-16, 4 figs. 

Reuss, a. E. 1845-1846. ' Die Versteinerungen der bohmischen Kreideformation : 1-58, pis. 

1-13 (1845) ; 1-148, pis. 14-51 (1846). Stuttgart. 
Saint-Seine, P. de & Casier, E. 1962. Poissons Fossiles des Couches de Stanleyville (Congo). 

Deuxieme partie. La Faune Marine des Calcaires de Songa. Ann. Mus. Afr. Cent. 8vo 

Sci. Geol., Tervuren, 44 : xi + 52 pp., 9 pis. 
Schaeffer, B. 1963. Cretaceous fishes from Bolivia, with Comments on Pristid Evolution. 

Amer. Mus. Novit., New York, 2159: 20 pp., 6 figs. 
Seilacher, a. 1943. Elasmobranchier-Reste aus dem oberen Muschelkalk und dem Keuper 

Wiirtembergs. Jb. Miner., Mh., Stuttgart (B) 1943 : 256-292, 50 figs. 
Smith, B. G. 1942. The Heterodontid Sharks. In The Bashford Dean Memorial Volume. 

Archaic Fishes : 647-770. Edit. Gudger, E. W. American Museum of Natural History, 

New York. 
Stensio, E. a. 1921. Triassic Fishes from Spitzbergen. Part /. 307 pp., 35 pis. Vienna. 



350 BRITISH WEALDEN SHARKS 

Stromer, E. 1927. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wiisten 

Agyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 9. Die 

Plagiostomen, mit Anhang iiber kano- und mesozoische Riickenflossenstacheln von Elasmo- 

branchiern. Abh. buyer. Akad. Wiss., Miinchen, 31, 5 : 1-64, pis. 1-3. 
Stubblefield, C. J. 1963. Summ. Progr. geol. Surv., Land. 1962 : 91 pp., London. 
WiLLisTON, S. W. 1900. Cretaceous Fishes. Selachians and Pycnodonts. Univ. geol. Surv. 

Kans., Topeka, 6 : 237-256, pis. 24-32. 
Woodward, A. Smith. 1887. On the Dentition and Affinities of the Selachian Genus Ptycho- 

dus Agassiz. Quart. J. Geol. Soc. Lond., 43 ; 121-131, pi. 10. 
1888. A Synopsis of the Vertebrate Fossils of the English Chalk. Proc. Geol. Ass., Lond., 

10 : 273-338, pi. I. 
1889. Catalogue of the Fossil Fishes in the British Museum [Natural History). 1. xlvii + 

474 pp., 17 pis. Brit. Mus. (Nat. Hist.), London. 
1891. The Hybodont and Cestraciont Sharks of the Cretaceous Period. Proc. Yorks. 

geol. [polyt.) Soc, Leeds, 12 : 62-68, pis. i, 2. 
■ 1904. On the Jaws of Ptychodus from the Chalk. Quart. J . Geol. Soc. Lond., 60 : 133-136, 

pi. 15. 
191 1. The Fossil Fishes of the English Chalk. Part VI. Mon. Palaeontogr. Soc, London, 

1910 : 185-224, pis. 39-46. 
■ 1912. The Fossil Fishes of the English Chalk. Part VII. Mon. Palaeontogr. Soc, 

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• 1916. The Fossil Fishes of the English Wealden and Purbeck Formations. Part I. 

Mon. Palaeontogr. Soc, London, 1916 ; 1-48, pis. i-io. 
1919. The Fossil Fishes of the English Wealden and Purbeck Formations. Part III. 

Mon. Palaeontogr. Soc, London, 1919 : 105-148, pis. 21-26. 
1932. Text-book of Palaeontology by K. A. von Zittel, 2, 2nd (English) edit, xvii + 464 pp., 

533 figs- London. 
Yabe, H. & Obata, T. 1930. On Some Fossil Fishes from the Cretaceous of Japan. Jap. J. 

Geol. Geogr., Tokyo, 8 : 1-7, pis. i, 2. 



PLATE I 

Fig. I. A block of bone-bed from the Weald Clay, Henfield Brick Co. pit, Heniield, Sussex. 
Teeth of Hyhodus basanns, H. brevicostahis and Hylaeobatis ornata are indicated by ' ba.', ' br.' 
and ' or.' respectively. P. 46920. Xi-5. 

Fig. 2. Caturus tenuidens Smith Woodward. A fragment of right dentary in lateral view. 
Weald Clay ; Henfield, Sussex. P. 46837. X4. 

Fig. 3. Hyhodus brevicostahis sp. nov. Upper anterior tooth in labial (a) and lingual (6) 
view. Wadhurst Clay ; Hastings, Sussex. P.11876. X3. 



Bull.B.M. [N.H.) Geol. 11, 7 



PLATE 1 







PLATE 2 
Hybodus brevicostatus sp. nov. 
Teeth of the holotype, P. 46973, Weald Clay ; Henfield, Sussex. 



All X4. 



Fig. I. Upper symphysial tooth in labial (a) and lingual yb) view. 

Fig. 2. Tooth from the fifth file of the right upper jaw in labial {a) and occlusal [b] view. 

Fig. 3. Tooth from the first file of the left lower jaw in labial [a) and occlusal (6) view. 

Fig. 4. Tooth from the sixth file of the left lower jaw in labial [a) and lingual [b) view. 

Fig. 5. Tooth from the eighth file of the left lower jaw in labial (a) and occlusal (6) view. 

Fig. 6. Tooth from the ninth file of the left upper jaw in labial [a] and lingual [b] view. 



Bull.B.M. {N.H.) Geol. 11, 7 



PLATE 2 






'W^A^'^' 



3b 





ft^yji^ 



mmmmym,, ,7,'^ 





PLATE 3 

Figs, i, 2. Hybodus brevicostatus sp. nov. Dorsal fin spines in posterior [a) and right lateral 
[b) view, natural size, with outline sections ( X 1-5) at the points marked. Fig. i, the holotype, 
P.46973, Weald Clay ; Henfield, Sussex. Fig. 2, P. 13268, Wealden Shales (overlying Hypsilo- 
phodon Bed) ; Cowleaze Chine, Isle of Wight. 

Fig. 3. Hybodus brevicostatus sp. nov. Thin section of a tooth from the holotype, P. 46973, 
Weald Clay ; Henfield, Sussex, x 12. 

Fig. 4. Lonchidion sp. Thin section of a dorsal fin spine, cut at the level marked in Text-fig. 
26A. P.47208, Weald Clay ; Henfield, Sussex, x 12. 



Biill. B.M. {N.H.) Geol. 11, 7 



PLATE 3 




PLATE 4 

Hylaeobatis ornata (Smith Woodward) 

Teeth from the Weald Clay of Henfield, Sussex. X5. 

Fig. I. Symphysial tooth in labial [a), occlusal [b], lingual (c) and lateral [d] view. P.47211. 

Fig. 2. Parasymphysial (first paired file) tooth in labial {a) and lingual [b] view. P. 47212. 

Fig. 3. Antero-lateral (second paired file) tooth in labial (a), occlusal [b), lingual (c) and 
lateral [d) view. P.47213. 

Fig. 4. Lateral tooth (probably fourth paired file) in labial {a), occlusal (6) and lingual (c) 
view. P.47214. 



Bull. B.M. (N.H.) Geol. 11, 7 



PLATE 4 







lb 



A"<'>,j\>"^"\'>\N 










PLATE 5 

Fig. I. Polyacrodus minimus {Aga.ssiz) . Vertical section of tooth crowii. P.47271, Rhaetic ; 
Holwell, Frome, Somerset, x 30. 

Fig. 2. Palaeobates angustissimus (Agassiz). Vertical section of tooth, P.47272, Mus- 
chelkalk ; Crailsheim, Germany, x 20. 

Fig. 3. Lonchidion breve breve sp. & ssp. nov. Vertical section of tooth crown cut through 
the labial process. P. 47275, Ashdown Beds, Cliff End bone-bed ; Cliff End, Sussex, x 50. 

Figs. 4, 5. Hylaeobatis ornata (Smith Woodward). Vertical sections of crowns of a para- 
symphysial tooth (Fig. 4, P.47276, x 20) and a posterior tooth (Fig. 5, P. 47277, X40), Weald 
Clay ; Henfiield, Sussex. 

Figs. 6, 7. Hylaeobatis ornata (Smith Woodward). Teeth in labial (a), occlusal (b) and 
lingual (c) view. Weald Clay ; Henfield, Sussex. Fig. 6. Posterior tooth (probably eighth 
paired file), P. 47216, x 10. Fig. 7. Postero-lateral tooth (probably sixth paired file), P. 47215, 
X5- 



Bull.B.M. {N.H.) Geol. 11, 7 



PLATE 5 




PRINTED IN GREAT BRITAIN 
BY ADLARD & SON LIMITED 
BARTHCLOMEW PRESS, DORKING 



ON CERTAIN TRIASSIC AND LIAS: 
REPRESENTATIVES OF THE FAMILY 
PHOLIDOPHORIDAE S. STR. 



O. NYBELIN 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

GEOLOGY Vol. ii No. 8 

LONDON: 1966 



2 a JAN 196 

ON CERTAIN TRIASSIC AND LIASSIC 

REPRESENTATIVES OF THE FAMILY 

PHOLIDOPHORIDAE S. STR. 



BY 



ORVAR NYBELIN . 

(Professor, Natural History Museum, Gothenburg, Sweden) 



Pp- 351-432 ; 15 Plates ; 16 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
GEOLOGY Vol. 11 No. 8 

LONDON: 1966 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily he completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 11, No. 8 of the Geological 
[Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 



Trustees of the British Museum (Natural History) 1966 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 21 January, 1966 Price £3 



ON CERTAIN TRIASSIC AND LIASSIC 

REPRESENTATIVES OF THE FAMILY 

PHOLIDOPHORIDAE S. STR. 



By ORVAR NYBELIN 



I. Introduction 



II. Systematic descriptions 

Genus Pholidophorus Agassiz 

Pholidophorus bechei Agassiz 

Pholidophorus latiusculus Agassiz . 

Pholidophorus (?) caffii Airaghi 

Pholidophorus cf. pusillus Agassiz . 
Genus Pholidolepis gen. nov. 

Pholidolepis dorsetensis gen. at sp. nov 
Genus Pholidophoroides Woodward 

Pholidophoroides crenulata (Egerton) 

Pholidophoroides limbata (Agassiz) . 
Genus Pholidophoropsis gen. nov. 

Pholidophoropsis caudalis (Woodward) 

Pholidophoropsis maculata sp. nov. 

III. Discussion ...... 

(a) The taxonomic relationship between the genera and species 
within the family Pholidophoridae s. str. and a preliminary 
diagnosis of the family ....... 

(b) Some phylogenetic aspects of the genera and species within 
the family Pholidophoridae s. str. ..... 

(c) Some brief comments on the derivation of the family 
Pholidophoridae s. str. ....... 

IV. Acknowledgements ........ 

V. References .......... 



Page 

354 
356 
356 

357 
368 

376 
382 

387 
387 
392 
393 
404 
411 
411 
416 

423 



423 
426 

429 
431 
431 



SYNOPSIS 

This paper is a revision, based mainly on the exoskeleton of the head and trunk, of a number 
of Upper Triassic and Liassic species formerly included in the genus Pholidophorus Agassiz. 
Special attention is paid to the shape of the preoperculum, the preopercular sensory canal and 
the squamation. Four species of Pholidophorus, including the type species. Ph. bechei, are 
redescribed. In the genus Pholidophoroides Woodward two species are redescribed. A new 
genus Pholidophoropsis is made for Pholidophorus caudalis Woodward and Pholidophoropsis 
maculata sp. nov. is described. A new genus and species, Pholidolepis dorsetensis, is made for 
part of the material previously included in Ph. caudalis. Relationships between the various 
species and genera of Pholidophoridae s. str., and between the Pholidophoridae and the Para- 
semionotidae and Leptolepidae are discussed. 

GEOL. 11,8 32 



354 CERTAIN TRIASSIC AND LIASSIC PH OLI DOPH ORI D AE 

I. INTRODUCTION 

The fundamental requisite for any discussion regarding the origin of the teleostean 
fishes is a knowledge of the species belonging to the family Pholidophoridae. Because 
of the vague definition of the genus Pholidophorus Agassiz 1832 many varied species 
have been attributed to it with the result that it has become one of the most extensive 
holostean genera, obviously containing many heterogeneous elements of dubious 
affinity. Consequently a revision of the genus Pholidophorus has long been highly 
desirable. 

In 1941 Woodward took up the question. Besides the genus Pholidophorus s. str., 
with Ph. bechei Agassiz as type species, he created three new genera : Pholidophoroides 
with Ph. crenulatus Egerton as type species and with Ph. caudalis Woodward as 
probably belonging to the same genus, Pholidophoristion with Ph. ovatus Agassiz 
as type species and Ph. micronyx Agassiz as second member, and Ichthyokentema for 
the two species Ph. purbeckensis Davies and Ph. brevis Davies. The last named genus 
has recently been treated in an excellent manner by Griffith & Patterson (1963), 
and its differences from Pholidophorus have proved to be so striking that a new 
family, Ichthyokentemidae, has been erected for it. The genera Pholidophoroides 
and Pholidophoristion have, however, not yet been thoroughly investigated and their 
validity consequently not proved. 

A rather large number of specimens referred to the species Ph. caudalis Woodward 
and belonging to the British Museum (Natural History) attracted my attention be- 
cause some of them showed a striking similarity to members of the genus Leptolepis. 
Because of this I have tried to study this material in more detail and have arrived 
at the conclusion that the specimens labelled as Ph. caudalis represent numerous 
different species. As Ph. caudalis was considered by Woodward as probably belong- 
ing to his new genus Pholidophoroides I have also found it necessary to take the type 
species Ph. crenulatus into consideration, and have tried to make a redescription of 
this species based on the excellent material in the British Museum (Natural History) . 
From Woodward's (1895) description of Pholidophorus limbatus Agassiz it seemed to 
me not unlikely that the species could have some relationship to the genus Pholido- 
phoroides. It is therefore included here. 

A definition of the genus Pholidophoroides required a comparison with the type 
species of the genus Pholidophorus. It is, however, not obvious which species 
should be considered as the type species of this genus. The genus Pholidophorus 
was erected by Agassiz (1832) for the two species Ph. latiusculus and Ph. pusillus 
from the Upper Trias of Seefeld, Tyrol. The diagnoses for the genus and for the two 
species are rather scanty and meaningless : " Pholidophorus Ag. Haringsgestalt. 
Grosse rautenformige Schuppen. Schwanzflosse ziemlich gleichlappig, indessen 
Ziehen sich die Schuppen noch an den obern Lappen hinauf. Riickenflosse den 
Bauchflossen gegeniiber. Afterflosse sehr klein. 

" I. Ph. latiusculus Ag. Grossere Schuppen. Im Verhaltniss breiter als der 
folgende. 

"2. Ph. pusillus Ag. Beide von Seefeld in Tyrol. In der Sammlung meines 
Freundes Dr. Alex. Braun, und letztere auch im Museiun in Carlsruhe." 



CERTAIN TRIASSIC AND LIASSIC PHOLI D OPHORID AE 355 

In his later work Agassiz (1833, 2 : 9) mentions the genus Pholidophorus for the 
first time, with the following five species : 

" I. Pholidophorus limbattis Agass. Ecailles frangees a leur bord posterieur. 
Corps tres-allonge. Lias : Lyme Regis. 

" 2. Pholidophorus dorsalis Agass. Caracterise par de longs chevrons sur le 
bord du premier rayon de la dorsale. Lias : Seefeld. 

" 3. Pholidophorus latiusculus Agass. Plus court ; ecailles plus grandes. See- 
feld. 

" 4. Pholidophorus pusillus Agass. Ecailles tres-petites. Seefeld. 

"5. Pholidophorus microps Agass. Tete petite ; ecailles en scie fine a leur bord 
posterieur, plus hautes que larges. Sohlenhofen." 

A more exhaustive description of the genus and its twenty species follows on p. 271 
of the same volume (1844) ; the first one treated is P/?. iec/zez Agassiz. Ph. latiusculus 
is only mentioned on p. 287 as the second species (after Ph. dorsalis) among those 
which were not figured for want of space but which the author intended to describe 
later on. The short note on Ph. latiusculus runs as follows : " Du lias de Seefeld et 
de Ljrme Regis. Espece tres-voisine de la precedente, mais plus petite, ayant la 
dorsale moins reculee ; elle n'a guere que deux a trois pouces de long." 

The first species ascribed to the genus Pholidophorus is thus Ph. latiuscidus, but 
Agassiz never named a type species of the genus nor a holotype of latiusculus, and 
never gave a figure of it ; the diagnoses cited above are too meagre and meaningless 
to allow an exact identification of the species. In his treatment of the fossil fishes 
from the Upper Trias of Seefeld, Kner (1866) tried to identify the three Pholidophorus 
species dorsalis, latiuscidus, and pusillus, but says regarding the proposed identifica- 
tion : " Ich hoffe hiedurch wenigstens anderen Palaontologen festere Anhaltspunkte 
zur Unterscheidung der Arten zu bieten und ihnen anschaulich zu machen, welche 
Formen mindestens mir den drei Arten von Agassiz zu entsprechen scheinen ; ob 
meine Deutung die richtige sei, dariiber mogen sie selbst dann entscheiden." Wood- 
ward (1895) obviously accepted the identification proposed by Kner and I cannot 
find any objection to this. 

Regarding the type species of the genus Pholidophorus Woodward did not choose 
the first named latiuscidus but hechei, the first species described and figured by 
Agassiz (1837, 2, pi. 39, figs. 1-4 ; 1844, 2 : 272). Woodward (1895 : 450-451) gives 
a good diagnosis of the latter species, which has more recently been investigated by 
Miss Rayner (1941, 1948), who was, however, principally interested in the study of 
the endocranium ; her description of the exoskeletal cranial bones and the accom- 
panying text-figures are thus rather schematic and do not allow a detailed comparison 
with other, closely related species. Consequently I have found it necessary to 
attempt a redescription and a new reconstruction of the exoskeletal cranial bones, 
based on some specimens belonging to the Department of Palaeontology, British 
Museiun (Natural History). 

The descriptions of Ph. latiusculus given by Kner (1866) and Woodward (1895) are 
also quite insufficient today. During a visit to Innsbruck in November 1963 I had 



356 CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHORID AE 

the opportunity to see in the Univ.-Institut fiir Geologie und Palaontologie some 
specimens from Seefeld, identified as Ph. latiusculus, among them a rather well 
preserved specimen figured by Kner (1866, pi. 3, fig. 3). As Ph. latiusculus seems to 
be closely related to Ph. hechei I find it convenient to append a short description of 
this species also, partly based on the specimens loaned from Innsbruck, partly on 
specimens belonging to the British Museum (Natural History), as well as of another 
interesting species, Pholidophorus caffii Airaghi, the holotype of which has been 
kindly placed at my disposal by the Director of the Museo Civico di Scienze Naturali 
" E. Caffi ", Bergamo, Italy. The description of a specimen probably belonging to 
Pholidophorus pusillus has also been included. 

Unless otherwise stated all the registered numbers given in the text refer to speci- 
mens in the British Museum (Natural History) collections. 



II. SYSTEMATIC DESCRIPTIONS 

Genus PHOLIDOPHORUS Agassiz 
1832 Pholidophorus Agassiz : 145. 

Preliminary diagnosis. Pholidophoridae of small to medium size. Exoskeletal 
cranial bones and scales with ganoin covering. Nasal well developed and well 
separated from its antimere by the frontals. Two well-developed supraorbitals. 
Maxillary not markedly stout and deep, posterior margin evenly rounded. Two 
supramaxillaries, overlapping dorsal margin of maxillary ; supramaxillary 2 without 
a marked process at antero-dorsal corner. Antorbital rather small ; five infra- 
orbitals. Preoperculum with preopercular sensory canal running nearer to anterior 
than to posterior margin. Lower jaw not markedly deep, its greatest depth being in 
the posterior third of its length ; dentary with a smooth dental part, separated from 
ornamented splenial part by strong ridge. Dorsal fin above base of ventral fins. 
Fulcra present along anterior margin of at least dorsal, pectoral and ventral fins ; 
caudal fin hemi-heterocercal with dorsal and ventral margins fulcrated. Scales 
rather thick with articulating pegs and with their posterior margin smooth ; anterior 
lateral line scales much deeper than broad. 

Type species. Pholidophorus hechei Agassiz. 

Remarks. The diagnosis given above for the genus Pholidophorus sensu stricto 
must at present be regarded as a preliminary one, and this mainly for two reasons. 
Firstly, it is based on two species only, the type species of the genus and Ph. latiusculus 
Agassiz ; only after a thorough study of the remaining species earlier ascribed to 
Pholidophorus or at least of a good nrunber of them, can the limits of this genus be 
made out and the common features of all its known species be established. Secondly, 
the present investigation only deals with external features, mainly the exoskeletal 
cranial bones ; an investigation of the endocranium and the visceral skeleton will 
undoubtedly reveal further characteristics of value for a definitive diagnosis. 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHORI D AE 357 

Pholidophorus bechei Agassiz 
(Pis. I, 2, 3 ; Text-figs, i, 2) 

1837 Pholidophorus bechei Agassiz, 2, pi. 39, figs. 1-4. 

1837 Pholidophorus onychius Agassiz, 2, pi. 39, figs. 5-7. 

1844 Pholidophorus bechei Agassiz, 2, i : 272. 

1844 Pholidophorus onychius Agassiz, 2, i : 274. 

1895 Pholidophorus bechei Agassiz ; Woodward : 450, pi. 12, figs, i, 2. 

1941 Pholidophorus Rayner : 230, text-fig. 10. 

1948 Pholidophorus bechei Agassiz ; Rayner : 318, pi. 21, fig. 45, text-figs. 24-29. 

1963 Pholidophorus bechei Agassiz ; Griffith & Patterson : 31. 

Preliminary diagnosis. Pholidophorus of medium size, up to about 200 mm. 
in total length. Nasal very large, anteriorly rounded and reaching beyond the 
anterior tip of the frontal. Posterior margin of preoperculum deeply notched. Pre- 
opercular sensory canal with about 17-19 tubules. 

HoLOTYPE. The first known specimen of this species is the one described and 
figured by de la Beche (1822) who, however, did not propose a scientific name for it ; 
according to Agassiz (1844, Vol. 2, pt. I : 273) this specimen was preserved in the 
collection of the Geological Society, London, but the specimen could not be found 
(March, 1965) in the collection of the Geological Survey Museum, London, where the 
British part of the Geological Society's collection is now housed. The specimen 
figured by Agassiz (1837, pl- 39- ^S- ^) belonged at that time to Miss Philpot, Ljone 
Regis. 

Material. The specimens used for the following description are Nos. 25276, 
38107, 38109, 39859, P. 154, P.1051, P.io52t/, P. 3586c, and P. 3589a, all belonging to 
the British Museum (Natural History), London. Unfortunately they are all more 
or less defective, especially in the ethmoidal region. 

Description. This species may attain a total length of about 200 mm. according 
to Woodward (1895 : 450), who also states that the length of the head with opercular 
apparatus is somewhat less than the maximum depth of the trunk and occupies 
one-fifth of the total length of the fish. As the main purpose of my investigation 
is to study the exoskeletal cranial bones of the type species of Pholidophorus, I have 
not examined the whole material in the British Museum (Natural History) but only 
some specimens showing these details. The largest of them, 25276, has a total length 
of about 164 mm., a standard length of about 140 mm. and a length of the head of 
about 37 mm., thus somewhat more than one-fifth of the total length [ca. 22-5%) 
and about one quarter of the standard length {ca. 26%). These values correspond 
rather well with those given by Woodward. Of the other larger specimens P.1051 
is defective but seems to be a little larger than 25276, while P. 1052^ is a little smaller 
than that specimen. 38107 and 38109 have a standard length of 113 mm. and about 
125 mm. respectively. The remaining three specimens are, however, much smaller 
with a standard length of about 80 mm. (P. 3589a), 76 mm. (P. 154), and 64 mm. 
(39859), respectively. 

Regarding the exoskeletal cranial bones no differences of taxonomic value can be 
observed between the larger and the smaller specimens, as far as their preservation 



358 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHO RI D AE 



allows a comparison. There exists, however, a marked difference regarding the 
development of the ganoin layer on the cranial bones as well as on the scales. In 
the largest specimens (PL 2, fig. 3) there is a continuous, thick layer of ganoin, 
more or less richly ornamented, on all exposed parts of the exoskeletal cranial bones 
and all the scales are thick. In the smaller specimens (64-80 mm. standard length ; 
PL 2, figs. I, 2) the ganoin covering is much thinner, on the exoskeletal cranial bones 
appearing as streaks, spots or small tuberculations, and the ganoin covering on the 
body scales seems to be remarkably thin in the anterior part of the body, gradually 
becoming thicker posteriorly. Even in 38107 (standard length 113 mm.) the scales 
on the anterior part of the body look thinner than in the posterior part. Perhaps 
this may be an indication of the beginning of the reduction of the ganoin covering 
which must have taken place in the phylogenetic development from the holostean 
to the teleostean stages of evolution. 

E,>'^ 55C 

I I 




poc orp 

Fig. I. Pholidophorus bechei Agassiz. Attempted restoration of head in lateral view, x 3.3. 
Ang, angular ; Ant, antorbital ; De. Spl, dentary ; Dpt, dermopterotic ; Dsph, dermo- 
sphenotic ; Ext, extrascapular ; Fr, frontal ; Ifo^-Ifo^, infraorbitals i to 5; lop, interoper- 
culum ; Mx, maxillary ; Na, nasal ; Op, operculum ; Pa, parietal ; Pmx, premaxillary ; 
Pop, preoperculum ; Ro, rostral ; Sbo, suborbital ; Smx-^, Smx^, anterior and posterior supra- 
maxillaries ; So-^, So^, anterior and posterior supraorbitals ; Sop, suboperculum ; Ssc, 
suprascapula ; ap, anterior pit-line ; Ac^, anterior division of supramaxillary pit-line ; ifc, 
infraorbital sensory canal ; ifc. com, ethmoidal commissure ; mp, middle pit-line ; orp, 
postmaxillary pit-line ; orpi, oral pit-line ; poc, preopercular sensory canal ; s. com, 
supratemporal commissure ; soc, supraorbital sensory canal. 



CERTAIN TRIASSIC AND LIASSIC PH OLI D OPHO RI D A E 359 

Exoskeletal skull roof 

The premaxillary {Pmx, PI. i ; PL 2, figs, i, 2 ; PI. 3, figs. 3, 4 ; Text-fig. i) has 
the form of an equilateral triangle with the dorsal margin almost straight, the postero- 
dorsal margin weakly S-shaped to fit the margin of the maxillary, and the antero- 
ventral margin a little convex and carrying a single row of 15 or more curved teeth. 
The slightly bulging outer surface carries a few ganoin tuberculations. 

The rostral {Ro, PL 2, figs. 2, 3 ; Text-figs, i, 2) is best preserved in the small 
specimen P.3589fl. Its median, bulging part is ornamented with a few ganoin 
spots, its ventro-lateral parts are, however, very defective and nothing can be made 
out with accuracy regarding the outline of the bone. In P. 1052^ parts of the rostral 
can also be observed. 

The nasal [Na, PL 2, figs. 1-3 ; Text-figs, i, 2) is comparatively very large. 
Anteriorly it is evenly rounded ; in its posterior part it tapers gradually backwards. 
In the smallest specimen investigated, 39859, its lateral margin has an almost semi- 
circular notch for the posterior nasal opening (PL 2, fig. i), in the largest specimen 
V.TO^zd, this opening is entirely surrounded by the bone (PL 2, fig. 2) just as figured 
by Miss Rayner (1941, text-fig. 10). In these two specimens as well as in P. 3589a 
the nasals of both sides are well separated by the anterior tip of the frontals ; nothing 
indicates that the nasals meet in the mid-line as figured by Miss Rayner. In the 
smallest specimen the dorsal surface of the nasal is quite smooth but in the two other 
specimens mentioned it is provided with a few ganoin spots, principally on the ridge 
indicating the course of the supraorbital sensory canal. 

The frontal {Fr, PL 2, figs. 1-3 ; Text-figs, i, 2) is pointed in its anterior part, 
mesial to the nasal, and from this region backwards it becomes progressively broader 
and has its broadest part posteriorly. Above the orbit the frontal margin is slightly 
concave, posterior to it the margin is almost straight and posteriorly directed. The 
postero-lateral comer of the frontal is rounded off and its posterior margin seems to 
be slightly wavy. The suture between the frontals of both sides is straight anteriorly 
but in the middle of its course it is irregularly sinuous, its path varying from specimen 
to specimen. The dorsal surface of the frontal is practically smooth in the smallest 
specimen (PL 2, fig. i) ; in P.3589a there are smaller and larger, partly confluent 
ganoin spots (PL 2, fig. 2) and in the large specimen, P.io52rf, the whole dorsal 
surface of the frontal is covered with ganoin forming radiating ridges of tubercula- 
tions on the lateral part of the anterior half of the bone (PL 2, fig. 3). 

There are two well-developed supraorbitals. The anterior one, supraorbital i 
{So-^, PL 2, figs. 1-3 ; Text-figs, i, 2), is situated in the angle between nasal and 
frontal ; its posterior half is rather broad and posteriorly rounded off, antero- 
laterally it tapers considerably. In the smallest specimen, 39859, its surface is 
smooth, in P. 3589a and especially in the large specimen, P.io52c^, the surface is 
ornamented with ganoin tuberculations. Posterior to supraorbital i there is an 
elongate supraorbital 2 in P.3589fl {S02, PL 2, fig. 3 ; Text -figs, i, 2) ; its mesial 
margin is slightly convex, fitting the concave margin of the frontal, anteriorly its 
margin is concave, fitting the posteriorly rounded supraorbital i and posteriorly it 
tapers gradually. Its dorso-lateral surface is covered with a rather thick ganoin 



36o 



CERTAIN TRIASSIC AND LIASSIC PH OLI DOPHORID AE 




Fig. 2, Pholidophorus bechei Agassiz. Attempted restoration of head in dorsal view. 
X 3.3. Lettering as in Fig. i. 

layer. In P.i052(f there is posterior to supraorbital i a small, rounded supraorbital 2; 
it seems not to be broken off posteriorly ; some bone fragments posterior to it may 
be other parts of the same bone, perhaps fragmented during an early stage in the 
ontogenetic development {So, PL 2, fig. 2). 

The dermosphenotic [Dsph, PI. i ; PL 2, fig. 2 ; Text-figs, i, 2) is missing in most 
specimens, and in T.io^2d only fragments of it are visible. In 38107, however, a 
rather well-preserved, roughly triangular dermosphenotic is present with its dorso- 
posterior margin convex and with its slightly concave and thickened anterior margin 
constituting the postero-dorsal border of the orbit. 

The shape of the exposed surface of the parietal (Pa, Text-figs, i, 2) is difficult to 
determine because of the poor preservation of this part of the skull roof, but it seems 
to be roughly square. 

The dermopterotic (Dpt, PL 2, figs. 1-3 ; Text-figs, i, 2) is comparatively narrow 
in its anterior part lateral to the posterior part of the frontal, but it broadens posterior 
to that bone on the dorsal surface of the skull, where it meets the parietal. 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHO RI D A E 361 

The extrascapular (Ext, PL i ; PL 2, fig. 2 ; Text-figs, i, 2) is heavily crushed in 
all specimens investigated by me, but it seems to have the usual triangular shape. 

Dermal bones of cheek and opercular apparatus 

The maxillary [Mx, PL i ; PI 2, figs, i, 2 ; PL 3, figs. 3, 4 ; Text-fig. i) is anteriorly 
pointed but it deepens ventrally, posterior to the premaxillary, so that a notch is 
formed for the premaxillary. The maxillary then becomes deeper with its greatest 
depth near its evenly rounded posterior margin ; its ventral margin is evenly convex. 
The dorsal margin, on the whole slightly concave, is overlapped by infraorbital i 
(lachrymal) and the two supramaxillaries ; anteriorly, at the deepening of the 
maxillary, its dorsal margin shows a shallow but marked concavity for the antero- 
ventral part of infraorbital i. The lateral surface of the maxillary is ornamented 
with a longitudinal striation which is parallel to the dorsal and ventral margins in 
the larger specimens ; however, in the smallest specimen, 39859, this ornamentation 
is only feebly marked (PL 2, fig. i), in the somewhat larger specimens, P. 154 and 
P.3589a, the ganoin streaks are broader and more irregularly arranged. Regarding 
the dentition along the ventral margin of the maxillary nothing can be said with 
accuracy because of the defective state of preservation ; in 38107 and 38109 the 
posterior part of the ventral margin shows, however, some markings which suggest a 
feeble dentition, and in P.3589fl the same part carries a delicate, comb-like dentition 
as in Leptolepis. 

The two supramaxillaries {Smx^, Smx^, PL i ; PL 2, fig. i ; PL 3, fig. 4 ; Text- 
fig, i) are situated dorsal to the posterior part of the maxillary, partly overlapping 
its dorsal margin. The anterior one, supramaxillary i, is comparatively small and 
roughly semicircular with its ventral margin straight. Its lateral surface has a 
ganoin ornamentation of irregular patches in the smaller specimens ; in the larger 
specimens they are confluent. In 38109 and P.1051 the anterior tip of the bone 
carries a well marked <-shaped crest parallel to its margins (PL 3, figs. 4, 5). The 
posteriorly situated supramaxillary 2 is much larger than supramaxillary i and 
almost triangular in shape with its greatest depth anteriorly ; its ventral margin is 
straight, its dorsal margin convex, and its anterior margin concave to fit the convex 
margin of supramaxillary i. Its antero-dorsal tip is slightly produced, and is 
apparently more pronounced in the smaller specimens. The lateral surface of 
supramaxillary 2 is ornamented with irregular ganoin spots in the smaller specimens; 
in the larger ones the entire surface is covered by a layer of ganoin with feebly marked 
tuberculations. 

The six bones of the infraorbital series (antorbital — five infraorbitals) are rather 
well preserved. 

The antorbital {Ant, PL 2, fig. i ; PL 3, fig. 2 ; Text-figs, i, 2), well exposed in 
25276, 39859 and P. 3586c, is comparatively small, almost triangular in shape with 
the angles rounded off. Its anterior and ventral margins are slightly convex and 
its posterior margin is straight or moderately concave. 

Infraorbital 1 (lachrymal) (Ifoi, PL 2, fig. i ; PL 3, figs. 3-5 ; Text-fig. i) is well 
preserved in 38109. It is elongate and deepest anteriorly. Its convex ventral 



362 CERTAIN TRIASSIC AND LIASSIC PH OLI DOPHORID AE 

margin has anteriorly a shallow notch, its postero-dorsal margin is not entirely 
visible but seems to be slightly concave. 

Infraorbital 2 {Ifo^, PI. i ; Text-fig. i), clearly visible in 38107, has the usual 
elongate shape with the dorsal margin slightly concave. The ventral margin is 
slightly convex. 

Infraorbital 3 (//03, PI. i ; PI. 2, fig. i ; Text-fig. i) is by far the largest bone in 
the series, stretching from the orbit to the anterior margin of the preoperculum, which 
it slightly overlaps. Its antero-dorsal and antero-ventral margins are concave, its 
dorsal margin is straight and its posterior margin is slightly convex. 

Infraorbitals 4 and 5 (//04, Ifo^, PI. i ; PL 2, fig. i ; Text -fig. i) are ahnost square 
with their anterior margins slightly concave and their posterior margins sUghtly 
convex. Infraorbital 5 is not entirely exposed in any of the specimens investigated 
but it seems to be a little smaller than infraorbital 4. 

A comparatively large suborbital {Sbo, PL i ; Text-fig. i) is situated posterior to 
infraorbitals 4 and 5 and seems to overlap the antero-dorsal part of the operculum. 
It is, however, generally crushed in the specimens available and in consequence of 
this, its outUne can only be made out in part. 

The preoperculum {Pop, PL i ; PL 2, figs, i, 2 ; PL 3, figs. 4, 5 ; Text-fig. i) has 
a very characteristic shape. The dorsal, almost vertical limb broadens continuously 
ventrally. The antero-ventrally directed limb is remarkably broad. The posterior 
margin of the bone is deeply notched below the angle between the two limbs, the 
postero-ventral comer projects backwards but is rounded off, not pointed as in Miss 
Rayner's reconstruction, the ventral margin is almost straight or slightly convex 
and the anterior margin shows a rather marked convexity below the middle of the 
bone. 

The operculum [Op, PL i ; PL 2, fig. i ; Text-fig. i) is roughly triangular ; the 
straight anterior, vertical margin and the straight postero-ventral margin meet at a 
low angle, the dorsal margin is evenly rounded. The anterior margin seems to be a 
little thickened. 

The suboperculum [Sop, PL i ; PL 3, figs. 4, 5 ; Text-fig. i) is comparatively 
large, but it is not entirely preserved in any of the specimens investigated and con- 
sequently its outline cannot be made out with accuracy. The antero-dorsal process 
anterior to the ventral tip of the operculum is comparatively strong. 

The posterior part of the inter operculum {lop, PL i ; PL 3, figs. 4, 5 ; Text-fig. i) is 
well exposed in 38107, 38109, and P.1051 ; its anterior part is not visible in any of 
the specimens investigated. 



Branchiostegal rays and gular plate 

Branchiostegal rays are exposed in 38107, 38109, and P.1051. In 38107 there are 
at least 12 branchiostegal rays on the right side {R. Br, PL i), in P.1051 at least 13 
rays may be distinguished, the six anterior ones still attached to the large ceratohyal 
2 {Ch^, PL 3, fig. 5), which is unfenestrated but provided with a deep groove for the 
hyoid artery. 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPH O RI D A E 363 

An elongate, anteriorly keeled, detached but not entirely exposed bone in P. 3589a 
may be interpreted as the gular plate {Gu, PI. 3, fig. i). A fragment of a gular plate 
is probably visible also in P. 154. 

Lower jaw 

The lower jaw is more or less exposed in most specimens ; in P.1051 it is almost 
entirely free from overlying bones (PL 3, fig. 5) ; it is a little defective but the outline 
of the defective parts can be followed through the impression in the rock. 

The dental and splenial parts of the dentary {De. Spl, PL i ; PL 2, figs, i, 2 ; PL 3, 
figs. 4, 5 ; Text-fig. i) are separated by a prominent ridge on the lateral surface of 
the bone ; the anterior part of its dorsal margin is only gently ascending ; at about 
the limit between the first and second third of the length of the lower jaw the dorsal 
margin of the dentary is abruptly thickened and ascends postero-dorsally. The 
boundary between the dentary and the angulo-supra-angular cannot be made out. 
The lateral surface of the splenial part is ornamented with thick striations and rugo- 
sities, whereas that of the dental part is quite smooth. In 38109 there are, as already 
stated by Miss Rayner (1948 : 319), some delicate teeth a little distance from the 
anterior tip of the dentary (PL 3, fig. 4) ; as far as I can see there are seven. 

The angulo-supra-angular {Ang, PL i ; PL 2, fig. i ; PL 3, figs. 4, 5) constitutes 
the posterior part of the lower jaw ; its depth equals about one-third of the length 
of the jaw. The lateral surface of the angulo-supra-angular has ventrally a con- 
spicuous ganoin ornamentation. 

No true dermarticular can be seen. The articular {Art, PL i ; PL 2, fig. i ; PL 3, 
figs. 4, 5) is partly visible in 38109. 

Sensory canal system of head 

The sensory canal system of the head is well developed, but because of the bad 
state of preservation of certain bones it cannot be described in every detail ; in 
particular the lack of a specimen with the bones of the snout in natural position 
makes the interpretation uncertain. 

The supraorbital sensory canal {soc, PL 2, fig. i ; Text-figs, i, 2) pierces the nasal, 
the frontal and the parietal. 

The nasal part of the canal is clearly visible in 39859, V.io$2d (PL 2, fig. 3), and 
P. 3589a (PL 2, fig. 2) ; it runs in a gentle arch from the antero-lateral to the postero- 
mesial margin of the bone, where it enters the frontal. In the large specimen, 
P.i052<f, there is a single pore visible on the dorso-mesial side of the canal a little 
anterior to the fenestra for the nostril. In the smaller specimen, P. 3589a, a corres- 
ponding pore may be traced on the dorso-lateral side of the canal ; in the smallest 
specimen no pores can be observed. 

The frontal part of the supraorbital sensory canal is very difficult to see because of 
the crushed state of the frontal in all specimens at my disposal. In the anterior 
part of the bone it runs, however, at first almost directly backwards, then it curves 
postero-laterally parallel to the lateral margin of the frontal and finally continues 
postero-mesially to the posterior margin of the bone. Pores or short tubules can 



364 CERTAIN TRIASSIC AND LIASSIC PHOLID OPH ORI D AE 

only be observed in P.io52i, and there the state of preservation does not allow any 
certain conclusions. According to Rayner (1948) there is great variability regarding 
the number and position of tubules and pores in this part of the supraorbital sensory 
canal ; my interpretation of the arrangement in P. 1052^ is given in Text-figs. 1,2. 
On the left side where the canal can be followed practically without interruption, 
except for its hindmost part, I have observed only a single short, laterally directed 
tubule in the posterior part of the anterior straight course of the canal ; at the curve 
there issue two somewhat longer, postero-mesially directed tubules from the mesial 
side of the canal and behind them two very short tubules on the same side of the 
canal ; on the postero-mesially directed part of the canal there are five very short 
tubules or pores. On the right side, where the canal is more defective, the tubules 
and pores seem to have on the whole the same arrangement, as far as they are 
discernible. 

The parietal part of the supraorbital sensory canal is visible on the left parietal ; 
the canal itself is very short, apparently without tubules or pores. It continues 
backwards as a well-marked groove, the anterior pit-line {ap, PL 2, fig. 2 ; Text-figs. 
1,2). 

The infraorbital sensory canals [ifc, PL 3, iig. 4 ; Text-figs, i, 2) of both sides are 
joined anteriorly by a slightly arched ethmoidal commissure in the rostral. In 
P. 1 05 2^ a short, posteriorly directed tubule can be seen on each side of the mid-line 
of the bone. 

The small antorbital contains in its ventral half the curved anterior part of the 
infraorbital sensory canal. From its convex dorsal side the canal gives off a short 
antorbital branch, ending with a comparatively large pore ; no more pores can be 
observed on this part of the canal except in 25276, where a pore seems to be present at 
the anterior angle between the canal and the antorbital branch (PL 3, fig. 2). 
According to Rayner (1941, text-fig. loB ; 1948, text-fig. 25) the antorbital branch 
joins the supraorbital sensory canal " to form a closed circuit around the eye ". 
I cannot, however, share this opinion. As already mentioned above no specimen 
seen by me has the exoskeletal bones of the snout preserved in natural articulation, 
but when trying to reconstruct this part of the skull I have not been able to arrive at 
a solution other than that given in Text-fig. i. Only much better preserved material 
regarding the mutual position of the bones in question can decide which solution is 
correct. 

Passing into infraorbital i (lachrymal) the infraorbital sensory canal pierces this 
bone on the whole parallel to its convex anterior and ventral margins. Judging 
from 38109 (PL 3, fig. 4) the canal gives off from its anterior side three short, antero- 
ventrally directed tubules and from its ventral side at least seven ventrally to postero- 
ventrally directed tubules, the middle ones being the largest ; as the bone is apparently 
not complete in its posteriormost part there might, consequently, exist one or more 
tubules in infraorbital i in addition to the ten observed by me. 

In infraorbital 2, only preserved in 38107, the sensory canal is not visible. 

In infraorbital 3 the sensory canal runs parallel to the concave anterior margin of 
the bone. In P. 154 it gives off two postero-ventrally directed tubules, the anterior 



CERTAIN TRIASSIC AND LIASSIC PH OLI D OPHO RI D AE 365 

one rather short, and two posteriorly directed tubules. In 38107 there seem to be 
only three tubules, only one of them posteriorly directed ; in the other specimens 
the state of preservation does not allow any observations regarding the tubules. 

In infraorbitals 4 and 5 the sensory canal gives off a single, posteriorly directed 
tubule in each bone. 

In the dermosphenotic, well preserved in 38107, the infraorbital sensory canal 
curves posteriorly ; from its convex dorsal side the canal gives off a rather wide, 
dorsally directed tubule, ending in a pore ; this pore is also visible on the defective 
dermosphenotic in P.io52i. 

From the dermosphenotic the canal enters the dermopterotic, piercing it along its 
lateral margin. To what extent the canal in the dermopterotic belongs to the sensory 
canal system of the head and to the cephalic division of the main lateral line, res- 
pectively, must be left unanswered. Judging from P.3589<* the posterior part of 
the canal gives off two dorsally directed short tubules, and the preopercular sensory 
canal seems to issue at the postero-lateral corner of the bone. 

Cephalic division of main lateral line 

A well-marked middle pit-line is visible in P. 1052^ on the left parietal posterior 
to the anterior pit-line ; it extends laterally over the broad posterior part of the 
dermopterotic. 

The cephalic division of the main lateral line passes over from the dermopterotic 
into the extrascapular, where it gives off mesially the supratemporal commissure, 
and continues posteriorly into the suprascapular. Because of the preservation of 
the extrascapular the canal and its tubules can be only partly observed ; in P.io52(^ 
two rather long tubules are given off from the lateral side of the canal, and at least 
four short tubules issue from the posterior side of the supratemporal commissure. 
In 38107 also, four short tubules are given off from the posterior side of the supra- 
temporal commissure decreasing in length mesially, but in this specimen only one 
large tubule can be observed with accuracy lateral to the main canal. 

The preopercttlar sensory canal runs on the whole parallel to the curved anterior 
margin of the preoperculum, about midway between its anterior and posterior mar- 
gins in the dorsal limb but decidedly nearer to its anterior margin in its antero- 
ventral limb. From its posterior side the canal gives off a series of tubules the number 
and arrangement of which are difficult to determine with precision in the largest 
specimens because of the thickness of the bone, especially in the dorsal limb of the 
bone ; in the small specimens the preoperculum is too defective to allow any detailed 
analysis of nmnber and position of the tubules, but their arrangement seems to be 
generally the same as in the larger specimens. In the an tero- ventral limb of the 
preoperculiuB there are in 25276, 38107, and 38109 thirteen postero-ventrally 
directed, curved tubules increasing in length from the anterior small one to the 
twelfth ; the thirteenth tubule is shorter and does not reach the postero-lateral 
margin of the bone. In P.1051 there is the same characteristic arrangement of 
the tubules, but their number cannot be counted with accuracy. Dorsal to this 



366 CERTAIN TRIASSIC AND LIASSIC PH OLI DOPHO RI D AE 

group of tubules the tubules are shorter and seem to vary a little in number and posi- 
tion. 25276 clearly shows six rather short tubules, in 38109 only five tubules can 
be observed, but the dorsalmost part of the bone is defective ; in 38107 and P. 154 
at least 4 short tubules are present, but also here their exact number is difficult to 
determine. In P.1051, in which the preoperculum is remarkably thick and, more- 
over, broken in its dorsal part, only three short tubules can be seen. The real number 
of small tubules in the dorsal part of the preoperculmn may be estimated as five or 
six and the total number of tubules belonging to the preopercular sensory canal may 
consequently be given as 18-19. 

On the lateral surface of the preoperculum at about its middle and anterior to 
the preopercular sensory canal two short grooves are clearly visible in most specimens, 
the dorsal one more or less horizontal, the ventral one on the whole vertical. It seems 
most likely that the horizontal groove represents the posterior portion of the anterior 
division of the supramaxillary line and the vertical one the post-maxillary line accord- 
ing to the nomenclature proposed by Stensio (1947). In P. 154, however, no supra- 
maxillary line can be observed on the preoperculum, but on infraorbital 3 an oblique 
groove is present, which may be interpreted as the anterior portion of the anterior 
division of the supramaxillary line. This difference in the development of the supra- 
maxillary line may merely be an individual variation rather than a feature of taxono- 
mic importance. 

The mandibular sensory canal and its tubules are not visible, only the openings 
of some separate tubules may be observed, but their total number cannot be deter- 
mined. 

In the largest specimen investigated, 25276, as well as in the small specimens 
39859 and P. 154, there is a small vertical groove on the lateral surface of the angular 
near its ventral margin ; this groove obviously represents the oral line according to 
Stensio (1947). 



Exoskeletal shoulder girdle and squamatien 

The suprascapula {Ssc, PI. i) is partly visible in 38107 and P.1052^, but in neither 
case can its outline be determined ; it is, however, a rather large bone. In 38107 a 
smaller, reniform bone postero-ventral to the suprascapula may be the supracleithrum, 
but no sensory canal can be observed in it. 

The cleithrum is only to a small extent exposed in the material studied by me and 
no description can be given. 

The scales are, according to Woodward (1895), arranged in approximately 40 
transverse rows ; in none of the specimens seen by me is the squamation uninterrupted 
but in 38107 there may be about 45 transverse rows reckoned from the posterior 
margin of the operculmn to the middle of the caudal fin. On the body there are 
about four longitudinal rows of scales which are deeper than broad, the lateral line 
scales being the deepest. All scales have an even posterior margin. 



CERTAIN TRIASSIC AND LIASSIC PH OLIDOPHORI D AE 367 

Lateral line 

The parts of the lateral line piercing the suprascapula and the supracleithrum 
are not visible in any of the specimens exhibiting these bones, and in the supra- 
cleithrum no tubules can be observed. In the suprascapula two rather wide tubules 
are clearly visible, the anterior one mesially, the posterior one postero-mesially 
directed. 

The lateral line runs almost straight along the sides of body and tail, ending at the 
base of the middle caudal rays. Each lateral line scale has a pore at about the middle 
of its lateral surface. 



Paired and unpaired fins 

The fins are as a rule more or less defective in all specimens investigated by me 
and consequently little can be added in this respect to the facts given by Woodward 
(1895 : 451). 

The pectoral fin seems to be of moderate size and with a moderate number of 
lepidotrichia, about 18 according to my counts, i.e. the same number as given by 
Woodward for the pectoral. The first lepidotrichium is rather stout and is provided 
with some fulcra. 

The ventral fin has a rather high number of lepidotrichia, about 14 or 15 (14 accord- 
ing to Woodward), the innermost ones rather delicate and branched almost to their 
bases. The first two lepidotrichia are comparatively short and undivided, the third 
one (the first branched lepidotrichium) carries some fulcra. 

The dorsal fin has, according to Woodward, about 12 lepidotrichia ; in my material 
no dorsal fin is complete. In P. 154 the anterior part of the fin is rather well preserved 
and shows anteriorly three undivided lepidotrichia followed by a long divided lepido- 
trichium with few but rather long fulcra. 

Regarding the anal fim Woodward gives no number of lepidotrichia, and in my 
material all anal fins are damaged and incomplete. On none of them have I observed 
any traces of fulcra, but as Woodward mentions the presence of fulcra on all fins, 
he must have seen some on the anal fin also. 

The caudal fin is hemi-heterocercal and provided with densely set fulcra on its 
dorsal as well as on its ventral margin. 

Remarks. Woodward (1895 : 450) considers Ph. onychius Agassiz to be identical 
with Ph. bechei. I have had no opportunity to see the holotype of Ph. onychius, 
belonging to the Oxford Museum, but 38109, originally labelled as Ph. onychius is, 
as far as I can see, in all respects a quite typical Ph. bechei, which supports Wood- 
ward's opinion. 

Horizon and locality. Lower Lias ; Lyme Regis, Dorset. 

GEOL II, 8 33 



368 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHO RID AE 

Pholidophorus latiusculus Agassiz 
(PI. 4 ; PL 5 ; PI. 15, figs, i, 2, 6, 7 ; Text-figs. 3, 4) 

1832 Pholidophorus latiusculus Agassiz : 145. 

1833 Pholidophorus latiusculus Agassiz ; Agassiz, 2 : 9. 

1844 Pholidophorus latiusculus Agassiz ; Agassiz, 2, i : 271, 287. 

1866 Pholidophorus latiusculus Agassiz ; Kner : 328, pi. 3, figs. 2, 3. 

1867 Pholidophorus latiusculus Agassiz ; Kner : 903, pi. 2, fig. i. 

1895 Pholidophorus latiusculus Agassiz ; (partim?) Woodward : 454, ? pi. 14, fig. 3. 

Preliminary diagnosis. Pholidophorus of small size, up to about 85 mm. in 
total length. Nasal not very large, anteriorly pointed. Posterior margin of pre- 
operculum with a shallow notch. Preopercular sensory canal with about 14-15 
tubules. 

Neotype. As already mentioned in the introduction, Agassiz did not name a 
holotype for this species. A species with this name is, however, described and 
figured by Kner (1866) ; the best preserved of his specimens, figured on PL 15, 
fig. 2, is still present in the collections of the Univ. Institut fiir Geologic und Palaon- 
tologie, Innsbruck, Austria, and may conveniently be chosen as neotype of Pholido- 
phorus latiusculus. The correctness of the identification is beyond doubt as can be 
seen by a comparison between the figure given by Kner (represented on PL 15, fig. i) 
and a photograph in natural size of the same specimen (PL 15, fig. 2). 

Material. The description given below is based on the following specimens : 
Three specimens, F.123 (neotype). Lit. F and No. 1028, all belonging to the Univ. 
Institut fiir Geologic und Palaontologie, Innsbruck, and two specimens, 33987 (in 
counterpart) and P. 1063, both belonging to the Department of Palaeontology, 
British Museum (Natural History) ; further, a rather disarticulated specimen, 
P.11780, showing a number of scales and some isolated cranial bones may also belong 
to this species or perhaps to Ph. cf. pusillus. 

Description. All the specimens, except P.11780, show the same characteristic 
shape, with the head and forepart of the body bent dorsally. The three Innsbruck 
specimens are of about the same size with a length without caudal (standard length) 
of ca. 75 mm. ; in the neotype the caudal fin is defective, in the two other specimens 
the caudal fin has a length of ca. 10 mm. ; the total length may consequently attain 
ca. 85 mm. Specimens 33987 and P.1063 are smaller ; the first with a total length of 
ca. 75 mm. and a standard length of ca. 65 mm., P. 1063 with a standard length of 
ca. 57 mm. The greatest depth of the body cannot be given as all specimens are 
more or less flattened through pressure. The length of the head seems to be roughly 
one quarter of the standard length. 

Exoskeletal skull roof 

The snout region is defective in all specimens available and the premaxillary and 
rostral bones are missing. 

The nasal {Na, PL 4 ; PL 5, fig. i ; Text-figs. 3, 4) is preserved only in the neotype 
and in P. 1063 ; it is comparatively large but its anterior part is pointed, not broadly 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHO RI D AE 



369 







De.Spl 



Fig. 3. Pholidophorus latiusculus Agassiz. Attempted restoration of head in lateral view. 
X 7 approx. 

Ang, angular ; De. Spl, dentary ; Dpt, dermopterotic ; Dsph, dermosphenotic ; Ext, extra- 
scapular ; Fr, frontal ; Ifo^-IfOr,, infraorbitals i to 5 ; lop, interoperculum ; Mx, maxillary ; 
Na, nasal ; Op, operculum ; Pa, parietal ; Pop, preoperculum ; Sbo, suborbital ; Smx^^, 
Smx^, anterior and posterior supramaxillaries ; So^, So^, anterior and posterior supraorbitals ; 
Sop, suboperculum ; ap, anterior pit-line ; hc^, anterior division of supramaxillary pit-line ; 
ifc, infraorbital sensory canal ; mp, middle pit-line ; orp, postmaxillary pit-line ; orp^, oral 
pit-line ; poc, preopercular sensory canal ; soc, supraorbital sensory canal. 

rounded as in Ph. bechei and the bone is not pierced by the posterior nasal opening. 
Its dorso-lateral surface is ornamented with irregular streaks and tuberculations. 

The frontal {Fr, PL 4 ; PL 5, figs, i, 2 ; Text-figs. 3, 4) has on the whole the same 
shape as in Ph. bechei, but its anterior part, mesial to the nasal, seems to be compara- 
tively more slender. As in that species the suture between the frontals of both sides 
is straight anteriorly but in the middle of its course somewhat sinuous ; in P. 1063 
(PL 5, fig. i) it curves at first a little to the left, then to the right and then again to 
the left. The entire dorsal surface of the frontal carries small ganoin rugosities 
which form low radiating ridges in its antero-lateral part. 

The two supraorbitals {So-^^, So^,, PL 4 ; PL 5, fig. i ; Text-figs. 3, 4) are of about the 
same size. The anterior one, supraorbital i, is situated posterior to the nasal ; in 
P. 1063 its anterior part is hidden below the posterior part of the nasal, but judging 
from the neotype its anterior part is rounded off, not drawn out into an antero-lateral 



370 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORIDAE 

projection as in Ph. hechei. The mesial margin of supraorbital 2 slightly overlaps 
the lateral margin of the frontal ; posteriorly its lateral margin is concave, fitting 
the antero-dorsal margin of the dermosphenotic. Both supraorbitals are ornamented 
with low ganoin ridges and rugosities. 

The dermosphenotic {Dsph, PL 5, fig. i ; Text-figs. 3, 4) is, judging from P. 1063, 
a rather small bone, pointed anteriorly and rounded posteriorly with its dorsal 
margin a little convex and its antero-ventral margin a little concave. 

The parietal {Pa, PL 5, fig. i ; Text-figs. 3, 4) and the dermopterotic {Dpt, PL 4 ; 
PL 5, figs. I, 2 ; Text-figs. 3, 4) cannot be described in detail because of the bad 
state of preservation which does not allow a determination of their boundaries. 

Only the lateral part of the extrascapular is preserved in P. 1063 [Ext, PL 5, fig. i). 
Apparently it is pushed over the posterior part of the dermopterotic, and conse- 
quently nothing can be made out with regard to its general shape. 



Dermal bones of cheek and opercular apparatus 

The maxillary {Mx, PL 4 ; PL 5, fig. i ; Text-fig. 3) is defective in all specimens 
but to judge from 33987 it is pointed anteriorly, rather as in Ph. bechei, and then 
deepens with its deepest part at its posterior rounded end. Its dorsal margin is 
slightly concave and is overlapped by infraorbital i (lachrymal) and the two supra- 
maxillaries. The lateral surface of the maxillary is provided with a dense longitudinal 
striation. Along the whole sUghtly convex ventral margin there is a feeble dentition 
like that in Leptolepis. 

The two supramaxillaries {Smx-^^, Smx^, PL 4 ; PL 5, fig. i ; Text-fig. 3) overlap 
the posterior part of the dorsal margin of the maxillary. The anterior one, supra- 
maxillary I, best preserved in the neotype (PL 4), is comparatively small, semicircular 
to almost triangular in shape ; its lateral surface is ornamented with well-marked 
concentric ridges or streaks. Among the few cranial bones preserved in P.11780 
there is a small semi-circular bone with a marked striation, which agrees rather well 
with supramaxillary i in the neotype. Supramaxillary 2 is fairly large and elongate 
with its dorsal and ventral margins slightly convex ; posteriorly it is rounded off ; 
its antero-ventral margin is concave, fitting the postero-dorsal margin of supra- 
maxillary I. The antero-dorsal tip of supramaxillary 2 is slightly thickened but 
not markedly produced. Its lateral surface is ornamented with numerous well- 
marked striations running parallel to its dorsal and ventral margins ; they are partly 
confluent in the middle of the bone. 

Unfortunately, of the bones of the infraorbital series, the antorbital is missing 
in all specimens investigated. 

Infraorbital i {lachrymal) {Ifo^, Text-fig. 3) is present only in 33987 and there only 
as a fragment of its anterior part ; nothing therefore can be made out with regard 
to its general shape. 

Infraorbital 2 {Ifo^, PL 5, fig. i ; Text-fig. 3) is preserved in P. 1063 ; it is, as usual, 
a small elongate bone which is situated below the orbit ; its posterior margin is 
postero-ventrally directed. 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHO RID AE 



371 



Dsph 




ifc 



Fig. 4. Pholidophorus latiusciilus Agassiz. 
view (possibly somewhat flattened). 



Attempted restoration of head in dorsal 
X 7 approx. Lettering as in Fig. 3. 



Infraorbital 3 (//03, PI. 4 ; PI. 5, figs. 1,3; Text-fig. 3) stretches from the orbit 
to the anterior margin of the preoperculum, which it overlaps as in Ph. bechei ; its 
general shape is also almost as in that species. 

Infraorbital 4 (i/04, PL 4 ; PI. 5, figs, i, 3 ; Text-fig. 3) has its anterior, dorsal 
and ventral margins almost straight whereas its posterior margin is shghtly convex ; 
it is about half as broad as infraorbital 3. 

Infraorbital 5 (//05, PI. 4 ; PI. 5, figs. I, 3 ; Text-fig. 3) can only be seen in part 
in the neotype, in P. 1063 and, in impression, in Innsbruck Lit. F ; it seems to have 
about the same general shape as infraorbital 4. 

The suborbital {Sbo, PL 4 ; PL 5, fig. i ; Text-fig. 3) seems to be a little broader 
than in Ph. bechei. Judging from the neotype its anterior margin lies almost vertical, 
whereas its posterior margin is slightly curved in the antero-ventral direction. 

The preoperculum [Pop, PL 4 ; PL 5, figs. 1,3; Text-fig. 3) is best preserved in 
the neotype, where it is partly hidden below infraorbital 3 and the suborbital, in 
P.1063 and, in impression, in Innsbruck Lit. F, where its postero-ventral part is 
lost. In 33987 it is rather defective. Its general shape resembles that of the pre- 
operculum in Ph. bechei, but its dorsal limb is broader and its postero-ventral part 



372 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPH ORI D AE 

does not protrude posteriorly as in that species ; the notch in its posterior margin is 
consequently much shallower. It is not possible to determine the course of the 
anterior margin of the preoperculum because of the overlying bones, but judging 
from the impression in Innsbruck Lit. F, the anterior margin has a slight convexity 
ventral to the middle of its length (PI. 5, fig. 3). In P.11780 there is a fragmentary 
bone which I interpret as the left preoperculum ; its anterior margin is rather well 
preserved and shows the same convexity as in the specimen just mentioned. 

The operculum {Op, PL 4 ; PI. 5, fig. i ; Text-fig. 3) has, on the whole, the same 
general shape as in Ph. hechei, but its dorsal part seems to be broader and its ventral 
tip is more obtuse ; its anterior margin is a little thickened except ventrally where 
the margin is slightly notched for the antero-dorsal process of the suboperculum. 

The suboperculum (Sop, PL 4 ; PL 5, fig. i ; Text -fig. 3) seems to be proportionately 
as large as in Ph. bechei but its posterior margin is defective in all specimens except 
P.11780, possibly belonging to this species ; in this specimen its posterior, well- 
preserved part shows a shallow concavity high on the posterior margin. 

Of the interoperculum [lop, PL 4 ; PL 5, fig. i ; Text-fig. 3) only the posterior 
part is visible in the neotype, in P. 1 1780 and, in impression, in Innsbruck Lit. F. 

Branchiostegal rays and gular plate 

The branchiostegal rays and the gular plate cannot be seen in any of the available 
specimens. 

Lower jaw 

The lower jaw is only partly exposed in the neotype and in 33987 and P. 1063 ; 
in Innsbruck Lit. F. the left lower jaw is exposed but very defective (PL 15, fig. 6) 
and in P.11780, possibly belonging to this species, an isolated right lower jaw is 
present but partly crushed (PL 15, fig. 7). These specimens suggest that the lower 
jaw in Ph. latiusculus has about the same shape as in Ph. hechei (PL 13, fig. 5). 

In all the specimens of Ph. latiusculus the ventral, splenial part of the dentary is 
separated from the dorsal, dental part by a well-marked ridge and its lateral surface 
is ornamented with thick ganoin rugosities and irregular striations. The dental 
part is anteriorly only gently ascending but further back it bends dorsally to form a 
small process, and then runs postero-dorsally ; its lateral surface is without orna- 
mentation. No teeth can be observed. 

The limit between the dentary and the angido-supra-angular [Ang, PL 15, fig. 7) 
is not clearly visible. The postero- ventral part of the lateral surface of the last named 
bone carries a well developed ornamentation. 

Sensory canal system of head 

The sensory canal system of the head seems to follow the same general pattern as 
in Ph. hechei, as far as the preservation of the material allows a comparison. 

The supraorhital sensory canal {soc, PL 5, fig. i ; Text-figs. 3, 4) pierces the nasal, 
the frontal and the parietal. 



CERTAIN TRIASSIC AND LIASSIC PH OLIDOPHORID AE 373 

In the nasal the canal runs almost straight from the antero-lateral to the postero- 
mesial part of the bone in P. 1063 ; no tubules or pores can be observed. In the 
neotype this part of the canal is not clearly visible. 

In the frontal the supraorbital sensory canal is rather well-marked as a slight ridge 
on the dorsal surface of the bone. Its course is on the whole the same as in Ph. 
bechei : an anterior part running almost straight posteriorly is followed by a curved 
part corresponding to the broadening of the frontal ; the posterior part runs at 
first in postero-mesial direction, then straight back to the middle of the posterior 
margin of the bone. P. 1063 has the following arrangement of tubules and pores 
(Text-figs. 3,4): from its anterior straight part the left canal gives off laterally two 
short tubules ; from its curved part three short tubules are given off from the mesial 
side ; the straight postero-mesially directed part has on its lateral side three very 
short tubules or pores ; the posteriormost part of the sensory canal is damaged and 
the presence of pores or tubules in that part cannot be made out. On the right side 
corresponding tubules are given off from the anterior and the curved parts of the 
canal, but further back only a single pore or tubule on the mesial side of the canal 
can be observed. In Innsbruck Lit. F. (PI. 5, fig. 2) there seem to be three laterally 
directed tubules issuing from the anterior straight part of the canal but only two 
pores from the curved part can be observed ; along the lateral side of the posterior, 
postero-mesially directed part of the canal there are four pores. 

The parietal part of the supraorbital sensory canal is well exposed in P. 1063 (PI. 5, 
fig. i). It is comparatively much longer than in Ph. bechei and on the right parietal 
there is a small shallow concavity on the dorsal surface of the canal which may be a 
pore, but its interpretation as a true pore is questionable. The sensory canal con- 
tinues posteriorly as a well-marked groove, the anterior pit-line [ap, PI. 5, fig. i ; 
Text-figs. 3, 4). 

The infraorbital sensory canal {ifc, Text-fig. 3) is quite unknown in its anterior 
parts because of the absence of the rostral and antorbitai bones. 

Only in 33987 is infraorbital i (lachrymal) present, and there only in its antero- 
ventral part ; it shows the anterior curved part of the canal in this bone with two 
antero-ventrally directed, pore-like tubules. For the rest the sensory canal and its 
tubules in infraorbital i are unknown. 

In infraorbital 2 the sensory canal is not visible. 

In infraorbital 3 the sensory canal pierces the bone parallel to its concave anterior 
margin. From its morphologically posterior side it gives off some tubules, the 
number of which is not easy to determine because of the thickness of the bone. In 
the neotype (PI. 4) there seem, however, to be three rather long posteriorly directed 
tubules situated close together ; antero-ventral to these tubules a short postero- 
ventrally directed tubule is present. In P. 1063 (PI. 5, fig. i) only two long, posteriorly 
directed tubules can be seen with certainty ; a short postero-ventrally directed tubule 
in the anterior part of the bone seems to be present but cannot be observed in detail. 

In infraorbital 4 the sensory canal gives off posteriorly a single tubule in the 
neotype. In infraorbital 5, which is only partly visible, no tubule can be observed 
with accuracy. 



374 CERTAIN TRIASSIC AND LIASSIC PHOLI D OPHORI D AE 

Judging from 33987, in which the posterior part of the dermosphenotic is preserved, 
the posteriorly curving sensory canal pierces this bone in its posterior part ; from 
the anterior side of the curve a rather wide tubule is given off in antero-dorsal 
direction. In P. 1063 the canal itself cannot be followed exactly, but the tubule is 
partly visible and seems to end in a pore at the dorsal margin of the bone. 

The sensory canal passes over into the dermopterotic and runs along its lateral 
margin. Judging from specimen Innsbruck Lit. F it seems to give off dorsally two 
short tubules. As in Ph. bechei the preopercular sensory canal seems to be given off 
near the postero-lateral corner of the bone. 



Cephalic division of main lateral line 

The middle pit-line {mp, PL 5, figs, i, 2 ; Text-figs. 3, 4) is well developed, stretch- 
ing from the postero-mesial part of the parietal to near the lateral margin of the 
dermopterotic. 

The cephalic division of the main lateral line passes over from the dermopterotic 
to the extrascapular, but as this bone is very badly preserved or missing nothing can 
be made out regarding this part of the canal and its tubules. 

The preopercular sensory canal runs, as in Ph. bechei, on the whole parallel to the 
anterior margin of the preoperculrmi. The tubules given off from its morpho- 
logically posterior side show the same general pattern as in Ph. bechei. In the 
neotype, the best preserved specimen in this respect, there seem to be three short, 
almost straight tubules dorsal to the angle on the posterior margin of the bone, the 
dorsahnost difficult to observe with accuracy as it is hidden below the suborbital, 
and ventral to the angle two shorter, slightly curved tubules ; these five dorsal 
tubules obviously correspond to the four to six shorter dorsal tubules in Ph. bechei. 
Ventral to these shorter tubules there is, as in Ph. bechei, a rather long tubule which, 
however, does not reach the posterior margin of the bone. Anteriorly it is followed 
by nine curved tubides which decrease in length antero-ventrally ; there are thus 
ten openings belonging to this group of tubules, but it seems as if the fourth tubule 
reckoned from the antero-ventral tip of the preoperculum is only a branch of the 
following tubule. The ten antero-ventral tubules in question obviously correspond 
to the thirteen antero-ventral tubules in Ph. bechei. In P. 1063 the shorter dorsal 
tubules are likewise five in number but the ventral part of the preoperculmn is too 
damaged to allow a count of the tubules. The preopercular impression in Innsbruck 
Lit. F shows five dorsally situated shorter tubules and ventral to them about nine 
tubules. The total ntmiber of tubules belonging to the preopercular sensory canal 
in Ph. latiusculus is consequently 14-15 against 17-19 in Ph. bechei. 

In P. 1063 (PI. 5, fig. i) the lateral surface of the preoperculiun shows one almost 
horizontal (hc^) and one vertical {orp) groove, obviously the posterior portion of the 
anterior division of the supramaxillary line and the postmaxillary line respectively 
as described above in Ph. bechei. An oblique, posteriorly directed groove on the 
lateral surface of infraorbital 3 may represent the anterior portion of the anterior 
division of the supramaxillary line. In the neotype (PI. 4) the postmaxillary line 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHORI D AE 375 

is hidden below infraorbital 3, but the other two lines are clearly visible, that on 
infraorbital 3, however, is not continuous but divided into two parts. In Innsbruck 
Lit. F. (PL 5, fig. 3) all the three lines mentioned can be traced on the impression. 
As compared with the corresponding lines in Ph. bechei the two lines on the pre- 
operculum are proportionately much longer and the line on infraorbital 3 seems to 
occur normally in all specimens. 

The mandibular sensory canal can partly be seen in the neotype and in P. 1063 ; 
the openings of separate tubules, marked by ganoin thickenings, are visible here and 
there ventral to the ridge separating the dental and splenial parts of the dentary ; 
their total number cannot be made out. 

In 33987, the only one in which the lateral surface of the angular is clearly exposed, 
a small vertical groove representing the oral line is visible on the angular near its 
ventral margin. 

Exoskeletal shoulder-girdle and squamation 

A suprascapula cannot be identified in any of the specimens investigated. In the 
neotype an almost triangular bone with strongly marked ganoin ridges on its lateral 
surface may be a supracleithrum {Scl, PI. 4). In the smaller specimen, 33987, the 
same bone is visible but the ganoin ridges are not as marked as in the neotype ; 
in P. 1063 (PI. 5, fig. i) a fragment of the same bone is present. In the last named 
specimen much of the cleithrum [CI, PL 5, fig. i) is exposed, and shows a large, deep 
notch above the insertion of the pectoral fin. 

The squamation (PL 4) is only partly preserved, but in the neotype about 38 
transverse scale rows can be observed, reckoned from the posterior margin of the 
operculum to the middle of the caudal fin base. The lateral line scales are the 
deepest, but also the rows immediately above and below the lateral line scales seem 
to be deeper than broad. All the scales are comparatively thick and have an even 
posterior margin. 

Lateral line 

Nothing can be made out regarding that part of the lateral line piercing the supra- 
scapula and the supracleithrum. The lateral line scales are arranged in an almost 
straight row along the side of the body {l.l, PL 4 ; cf. also PL 15, fig. 2). 

Paired and unpaired fins 

Very little can be said regarding the fins in this species. 

The pectoral fin is partly preserved in all specimens investigated but only in P. 1063 
can the number of lepidotrichia be counted with accuracy (PL 5, fig. i) ; there seem 
to be about 19 lepidotrichia, the first one carrying some small fulcra. 

Only a few lepidotrichia belonging to the ventral fin are preserved in specimen 
33987, in others the ventral fin is missing. 

The dorsal and anal fins are lacking in all specimens seen by me, but according to 
Kner (1867, pi. 2, fig. i) the dorsal fin is situated approximately above the ventral 
fins as in Ph. bechei. 



376 CERTAIN TRIASSIC AND LIASSIC PH OLI DOPHO RID AE 

The caudal fin is hemi-heterocercal and carries well-developed fulcra on its dorsal 
and ventral margins. 

Remarks. Apart from small differences, mainly in the ornamentation of the 
exoskeletal cranial bones, all the specimens mentioned above except P. 1 1780 are so 
similar that they may be considered to belong to one and the same species. Whether 
the specimens from Italian localities attributed to Ph. latiiisculns by authors such as 
de Alessandri, Bassani, Costa, Mariani etc., are identical with the specimens described 
above is a question which at present must be left open, as I have had no opportunity 
to see them. 

Horizon and locality. Upper Trias ; Seefeld, Tyrol, Austria. 

Pholidophorus (?) caffii Airaghi 

(PL 6, fig. I ; Text-fig. 5) 

1908 Pholidophorus Caffii Airaghi : 3, text-fig. 2. 

1914 Pholidophorus latiusculus Agassiz ; (partim) Bassani : 379. 

1920 Pholidophorus latiusculus Agassiz ; (partim) Alessandri : 96. 

1937 Pholidophorus latiusculus Agassiz ; Boni : 132, pL 4, fig. 4, pi. 5, fig. 3, text-figs. 9, 10. 

Diagnosis. Pholidophorus (?) of small size, up to 58 mm. in total length. Greatest 
depth of body about one quarter of the standard length or slightly more. Depth of 
caudal peduncle about one-tenth of the standard length. Maxillary and lower jaw 
rather short. Infraorbitals 3 and 4 deeper than broad. Preoperculum broad, its 
posterior margin without a marked notch. Preopercular sensory canal running 
nearer to posterior than to anterior margin of bone in dorsal limb, in ventral limb 
at about the middle, and with about eight tubules. Scales thick, not serrated 
posteriorly, arranged in about 35 transverse rows counted from posterior margin of 
operculum to middle of caudal fin base. A few longitudinal rows of body scales 
deeper than broad. 

HoLOTYPE. Specimen originally described by Airaghi (1908) and belonging to the 
Museo Civico di Scienze Naturale di Bergamo, Italy. The only specimen. 

Description. The total length of the holotype is 58 mm., the length from the 
tip of the snout to the hindmost lateral line scale (standard length) is 49 mm. The 
estimated depth of the body is slightly more than one quarter (26-5%) of the standard 
length and the depth of the caudal peduncle is about one-tenth of that length. 
The length of the head is estimated to be slightly less than one quarter of the standard 
length. The maxillary and lower jaw are, comparatively, a little shorter than in 
Ph. bechei and Ph. latiusculus. 

The position of the paired and unpaired fins is clearly visible. The base of the 
ventral fin is situated a little nearer to the hindmost lateral line scale than to the tip 
of the snout, and the distance between the base of the ventral and the origin of the 
anal is about 75% of the distance between the bases of the pectoral and ventral 
fins. The dorsal fin begins above the ventral fin base. A good figure of the entire 
fish is given by Boni (1937, pi. 5, fig. 3). 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPH O RI D AE 



377 



The cranial bones are, unfortunately, partly defective ; most bones belonging to 
the exoskeletal skull roof are lacking and many of the other cranial bones are poorly 
preserved. Consequently the following description is rather incomplete. 

Exoskeletal skull roof 

The premaxillary, rostral, nasal, frontal, supraorbital, dermosphenotic and 
dermopterotic bones are missing or represented by insignificant fragments. Parts 
of the parietal and the extrascapular seem to be present {Pal , Ext?, PL 6, iig. i), but 
their shape cannot be made out. 

Dermal bones of cheek and opercular apparatus 

Only the posteriorly directed lateral part of the comparatively short maxillary is 
present {Mx + Smxi, PL 6, fig. i ; Text-fig. 5). It is, however, very defective, 
almost the entire ventral part is missing and consequently nothing can be made out 




Fig. 5. Pholidophorns (?) caffii Airaghi. Holotype. Attempted restoration of head in lateral 
view. X 10-5. 

Ang, angular ; Ant, antorbital ; De. Spl, dentary ; IfOj-Ifo^, infraorbitals i to 5 ; lop, 
interoperculum ; Mx + Smx^, maxillary and anterior supramaxillary ; Op, operculum ; 
Pop, preoperculum ; Sbo, suborbital ; Smx^, posterior supramaxillary ; Sop, suboperculum ; 
Acj, anterior division of supramaxillary pit-line ; ifc, infraorbital sensory canal ; mdc, mandibular 
sensory canal ; orp, postmaxillary pit-line ; orp^, oral pit-line ; poc, preopercular sensory canal. 



378 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORIDAE 

regarding the depth of the maxillary or of the dentition. The lateral surface of the 
preserved part is, however, ornamented with longitudinal streaks and, posteriorly, 
with some short oblique ridges. The posterior end of the maxillary is broken off, 
and thus its shape, if rounded or notched, cannot be stated. According to Boni 
(1937, text-fig. 9) the posterior part of the maxillary is wedged between infraorbital 3 
and the antero-ventral limb of the preoperculum, but the part in question is, as far 
as I can see, the antero-dorsal part of the ventral limb of the preoperculum, and not 
part of the maxillary. 

Boni's text-fig. 9 shows only a single supramaxillary , obviously corresponding to 
supramaxillary 2 in other species ; anterior to it the figure shows, however, a roughly 
triangular or semicircular elevation on the dorsal margin of the maxillary, corres- 
ponding to supramaxillary i in other forms. At first I was inclined to consider 
this eminence as not belonging to that bone, but as soon as I had the opportunity of 
examining the holotype I could confirm that there was no division between this 
elevation and the rest of the maxillary. Whether this is a secondary, perhaps 
individual, fusion of the two bones or a primitive stage of development, characteristic 
of Ph. caffii, is a question which must be left open until more specimens of this species 
are available for study. Supramaxillary 2 {Smx^, PL 6, fig. i ; Text-fig. 5) is rather 
well preserved ; it is almost rectangular, its antero-dorsal corner forms a blunt 
short process, and its lateral surface is ornamented with rather dense, horizontal 
striations. 

All the bones of the infraorbital series, an antorbital and five infraorbitals, are 
present. 

The antorbital {Ant, PL 6, fig. i ; Text-fig. 5) is rather well preserved but its outline 
cannot be made out in detail ; it is large in comparison with the antorbital in Ph. 
bechei and tapers postero-dorsally. 

Infraorbital i {lachrymal) {Ifo^^, PL 6, fig. i ; Text-fig. 5) was exposed by prepara- 
tion ; it is rounded anteriorly, its ventral margin is convex, its dorsal margin concave 
and the exposed dorso-lateral surface is slightly concave. 

Infraorbital 2 (//og, PL 6, fig. i ; Text -fig. 5) is well preserved but broken into two 
pieces ; it has a generally elongate shape with the dorsal margin slightly concave, 
the ventral margin slightly convex. 

Infraorbital 3 (//og, PL 6, fig. i ; Text-fig. 5) is likewise well preserved except 
that its postero-dorsal corner is broken off ; it is markedly deeper than broad and 
its dorsal margin constitutes an obtuse angle so as to fit the ventral margins of 
infraorbital 4 and the suborbital. 

Infraorbital 4 {Ifo^, PL 6, fig. i ; Text -fig. 5) is partly incomplete but seems to be a 
little deeper than broad. 

Infraorbital 5 (//05, PI. 6, fig. i ; Text -fig. 5) is better preserved and seems to be 
almost square and only a little smaller than infraorbital 4. 

Posterior to infraorbitals 4 and 5, the ventral part of the suborbital {Sbo, PL 6, 
fig. I ; Text -fig. 5) is clearly visible but it is broken into two pieces on to the lateral 
crest of the hyomandibular ; regarding its outline it can only be said that its posterior 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHO RI D AE 379 

margin seems to be straight and vertical, thus probably not overlapping the anterior 
margin of the operculum, and that its postero- ventral margin is a little convex. 

The preoperculum {Pop, PL 6, fig. i ; Text-fig. 5) is, comparatively, very broad, 
especially in its antero-ventral limb, the antero-dorsal margin of which shows a 
rather pronounced convexity ; the postero-ventral margin of the preoperculum is 
damaged but it seems to have been almost straight or only shghtly concave and thus 
without the notch characteristic of the preoperculum in Ph. latiusculus and especially 
in Ph. bechei. The uppermost end of the preoperculum is defective. 

The operculum {Op, PI. 6, fig. i ; Text-fig. 5) seems to have the usual almost 
triangular shape, but as its anterior and dorsal margins are damaged its shape cannot 
be made out exactly. Its ventral tip is rounded, about as in Ph. latiusculus, and as 
in that species the anterior margin of the operculum is slightly notched ventraily to 
receive the antero-dorsal process of the suboperculum. 

The suboperculum {Sop, PI. 6, fig. i ; Text-fig. 5) is only partly preserved ; its 
antero-dorsal process is well developed. 

The inter operculum {lop, PI. 6, fig. i ; Text-fig. 5) seems to be roughly triangular 
with the posterior margin slightly convex ; its lateral surface carries a few striations 
parallel to its margins. 

Branchiostegal rays and gular plate 

The posteriorly situated, rather short but broad branchiostegal rays {R. Br, PL 6, 
fig. i) are visible ventral to the preoperculum and the interoperculum. 
The gular plate is not visible. 

Lower jaw 

The lower jaw is, like the maxillary, comparatively short. Anteriorly the dorsal 
margin of the dental part of the dentary {De. Spl, PL 6, fig. i ; Text-fig. 5) is only 
gently ascending, but as most of the dorsal margin of the lower jaw is covered by 
overlying bones, its outline cannot be followed. No teeth could be observed on the 
exposed dorsal margin of the dentary. The lateral surface of the dental part is 
smooth and that of the splenial part is only feebly ornamented but shows clearly 
the tubules of the mandibular sensory canal ; the ridge separating the two parts of 
the dentary seems to be comparatively weak. 

The ventral part of the angular {Ang, PL 6, fig. i ; Text-fig. 5) is well exposed ; 
its lateral surface is practically smooth. 

Sensory canal system of head 

Regarding the sensory canal system of the head, almost nothing of the supra- 
orbital sensory canal can be determined because the nasal and frontal bones are 
lacking. On the fragment thought to belong to the parietal a single short canal is 
visible. This might be interpreted as the posteriormost part of the supraorbital 
sensory canal, but as it is not followed by a pit-line this interpretation is rather 
uncertain. No canals, tubules or pores can be observed in the bone fragment con- 
sidered as part of the extrascapular. 



38o CERTAIN TRIASSIC AND LIASSIC PH O LI DOPHO RI D AE 

The infraorbital sensory canal [ifc, Text-fig. 5) can be followed for almost its entire 
length ; only the ethmoidal commissure and the part in the dermosphenotic and 
dermopterotic are missing. In the antorbital the canal describes an arch in its 
ventral part and gives off postero-dorsally an antorbital branch ending in a pore ; 
at the point where the antorbital branch arises there seems to be a rather large pore. 
In infraorbital i (lachrymal) the sensory canal can be followed for most of its length 
but its state of preservation allows no statements regarding the number or disposition 
of its tubules. In infraorbital 2 the sensory canal is partly visible ; an opening at 
the posterior end of the bone is not a real pore but an artifact due to damage. In 
infraorbital 3 the sensory canal gives off three postero-ventrally directed and short 
but strongly marked and widely separated tubules, the antero-ventral one being the 
shortest, the dorsal one the longest. The anterior part of infraorbital 4 is obviously 
pierced by the sensory canal, but no tubule can be seen with certainty. In infra- 
orbital 5 the sensory canal can be traced but no tubule is visible. 

Cephalic division of main lateral line 

The preoperctdar sensory canal seems to pierce the dorsal limb of the preoperculum 
nearer to its posterior than to its anterior margin ; in the ventral limb it runs at 
about the middle of its length, not nearer to its anterior margin as in Ph. bechei and 
Ph. latiusculus. The tubules given off from the morphologically posterior side of 
the canal are short but rather wide. Only eight tubules seem to be present, two in 
the dorsal limb of the preoperculum dorsal to and at the posterior angle of the bone 
respectively, one short tubule farther ventrally and five tubules in the ventral part 
of the ventral limb of the bone. Consequently, the general arrangement of the 
tubules is not unlike that in Ph. bechei and Ph. latiusculus , but the tubules in the 
ventral group are not long and narrow as in the two former species and the total 
number of tubules is only about 8 as against 17-19 in Ph. bechei and 14-15 in Ph. 
latiusculus. 

On the lateral surface of the preoperculiun there are two grooves, one horizontal 
and one vertical, (hc-^, orp, PL 6, fig. i ; Text-fig. 5), meeting at a right angle ; the 
horizontal groove represents the posterior portion of the anterior division of the 
supramaxillary line and the vertical one the postmaxillary line according to the 
nomenclature proposed by Stensio (1947). 

The mandibular sensory canal {mdc, PL 6, fig. i ; Text -fig. 5) is exposed to a small 
extent in the anterior, broken portion of the splenial part of the dentary ; it is 
impossible to say how many tubules the canal gives off in that part of the bone, but 
posterior to the exposed part of the canal five short tubules are visible. Near the 
ventral margin of the angular there is a pore belonging to that part of the mandibular 
sensory canal which pierces the angular, and dorsal to this pore there is a short, 
vertical groove, representing the oral line {orp^, PL 6, fig. i ; Text-fig. 5). 

Exoskeletal shoulder-girdle and squaniation 

Dorsal to the operculum there is a large suprascapula [Ssc, PL 6, fig. i) but it is 
crushed and partly defective and consequently its outhne cannot be determined 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHO RI D AE 381 

exactly. Postero-ventral to it there is a suprachithrum {Scl, PI. 6, fig. i) of about the 
same size and shape as the suprascapula, but this bone also is crushed and defective. 
Almost the entire cleithriim is missing but its impression suggests a rather robust 
bone. Posterior to this impression there is a deep bony plate which may be inter- 
preted as a postcleithrum . 

The scales are arranged in about 35 transverse rows counted from the posterior 
margin of the supracleithrum to the middle of the caudal fin base (= lateral line 
scales). On the body the lateral line scales and those in the longitudinal rows 
immediately dorsal and ventral to them, three rows in all, are deeper than broad. 
All the scales are comparatively thick and, as far as can be judged, with a regular 
posterior margin. 

Lateral line 

Those parts of the lateral line piercing the suprascapula and the supracleithrum 
are not visible ; in the suprascapula, however, a short but rather wide, mesially 
directed tubule can be seen ; in the supracleithrum no tubules are visible. 

The lateral line scales {l.l, PI. 6, fig. i) are arranged in a straight row from the 
second scale to the last one at the base of the middle caudal rays. 

Paired and unpaired fins 

All fins are present but at least in part more or less defective. 

The pectoral fin is of moderate size. Boni (1937) gives the number of its lepido- 
trichia as 11, a remarkably low number. There are, indeed, 11 rather stout lepido- 
trichia clearly exposed, but ventral to the postcleithrum and easily overlooked 
there is a cluster of broken, more slender lepidotrichia ; these may represent the 
posterior rays of the pectoral fin and their number may be at least 7-8, and conse- 
quently the total number of pectoral lepidotrichia may be estimated as about 18 
or about the same number as in Ph. bechei and Ph. latiusculus. The first lepido- 
trichium carries some very small fulcra (Fu, PL 6, fig. i). 

The ventral fin is rather defective. For this fin Boni gives the number of lepido- 
trichia as 5, which is too low ; as far as I can see there are at least two small, un- 
divided lepidotrichia anteriorly, followed by 6 divided and branched lepidotrichia, 
the anterior one fulcrated. Posterior to these lepidotrichia the fin is damaged and 
the total number of ventral fin lepidotrichia thus remains unknown. 

The dorsal fim is well preserved but the number of lepidotrichia is difficult to 
interpret. Anteriorly there are two undivided lepidotrichia, the anterior one 
rather short and almost scale-like ; there follows a third, rather long, apparently 
undivided lepidotrichium with a comparatively long and slender rod (fulcrum?) 
in front of it ; posterior to these three there are 8 divided and branched lepidotrichia, 
the anteriormost of which is richly fulcrated. Consequently, the total number of 
dorsal fin lepidotrichia is at least 11 as compared with g (10?) as stated by Boni. 

The anal fin is not as well preserved as the dorsal fin and the number of its lepido- 
trichia cannot be stated with accuracv. There are, however, two short, undivided 



382 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORIDAE 

lepidotrichia anteriorly followed by at least 7 divided and branched ones, the 
anteriormost of which with a few, long fulcra ; thus there are at least g lepidotrichia 
as against the 5-6 as stated by Boni. 

The caudal fin is hemi-heterocercal and its dorsal as well as its ventral margin is 
densely set with fulcra. 

Remarks. Pholidophorus cafiii, described as a new species by Airaghi (1908), 
has been considered merely a synonym of Ph. latiuscidus by Bassani (1914), de 
Alessandri (1920) and Boni (1937). It is, however, beyond doubt that the specimen 
on which Airaghi founded his species, redescribed above, does represent a species 
clearly distinguishable from Ph. latiusculus in the sense of Kner. Differences in the 
relative length of maxillary and lower jaw, the general shape of the preoperculum, 
the course of the preopercular sensory canal and the number of its tubules clearly 
separate Airaghi's Pholidophorus caffii from Kner's Ph. latiusculus. In my opinion 
differences in the preoperculum and the course of the preopercular sensory canal 
are of such a degree that the placing of caffii in the genus Pholidophorus s. str.may 
even be questionable. This problem will be discussed in more detail later on (p. 423). 

Horizon and locality. Upper Trias ; Viciarola, near S. Pellegrino (Val Brem- 
bana), Italy. 

Pholidophorus of. pusillus Agassiz 
(PL 6, fig. 2 ; Text-fig. 6) 
1895 Pholidophorus latiusculus Agassiz ; Woodward : 455 (partim). 

Preliminary diagnosis. Pholidophorus of small size, up to 65 mm. in total 
length, similar to Ph. (?) caffn but differing from that species i.a. in the following 
features : Maxillary and lower jaw not markedly short. Infraorbitals 3 and 4 
considerably broader than deep. Preoperculum similar to that of caffii but pre- 
opercular sensory canal with about 11 tubules, longer than those in caffi. Scales 
arranged in about 38-40 transverse rows counted from posterior margin of operculum 
to middle of caudal fin base. 

Material. Specimen No. P. 4418 in the Department of Palaeontology, British 
Museum (Natural History). 

Description. The total length of the specimen is about 65 mm., the standard 
length is estimated as 56 mm. The greatest depth of the body cannot be deter- 
mined. The depth of the caudal peduncle is about one-tenth of the standard length. 
The length of the head is estimated to be shghtly less than one quarter of the standard 
length. The maxillary and lower jaw are relatively longer than in Ph. bechei and 
Ph. latiusculus, and even more so than in Ph. (?) caffii. 

The fins are not preserved except for some lepidotrichia of the anal fin and the 
entire caudal fin ; the position of the dorsal fin in relation to the ventral fins is thus 
unknown. 

The cranial bones are very defective ; all bones belonging to the exoskeletal 
skull roof are lacking and many of the other cranial bones are fragmentary. Thus 
only the following rather incomplete description can be given. 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHORI D AE 



383 



Dermal bones of cheek and opercular apparatus 

Only the posteriorly directed part of the comparatively long maxillary {Mx, PI. 6, 
fig. 2 ; Text-fig. 6) is preserved ; apart from its relatively greater length it seems to 
have the same general shape as in Ph. bechei and Ph. latiusculus, but its ventral 
margin is defective and consequently nothing can be said about its real depth or 
about the dentition. Neither can it be determined with any accuracy whether its 
posterior margin is rounded or slightly notched ; judging from an impression on 
the preoperculum behind the broken end of the maxillary it looks, however, as if its 
posterior margin was slightly notched. The lateral surface of the maxillary is 
richly ornamented with ganoin spots and irregular longitudinal streaks and ridges 
except along its postero-dorsal margin, indicating that supramaxillary 2 may overlap 
the maxillary a little. 

Of the supramaxillaries, supramaxillary i {Smx-^, PL 6, fig. 2 ; Text-fig. 6) is 
almost semicircular ; it is well separated from the maxillary, but seems not to overlap 
the dorsal margin of that bone ; its lateral surface carries a striation approximately 
parallel to the convex dorsal margin of the bone, similar to the striation on the dorsal 




Fig. 6. Pholidophorns cf. pusillus Agassiz. Attempted restoration of head in lateral view. 
B.M. No. P.4418. X 10. 

Ang, angular ; De. Spl, dentary ; //oi, //03, //04, infraorbitals i and 3-4 ; lop, interoperculum ; 
Mx, maxillary ; Op, operculum ; Pop, preoperculum ; Sbo, suborbital ; Smx^, Smx^, anterior 
and posterior supramaxillaries ; Sop, suboperculum ; poc, preopercular sensory canal. 

GEGL. II, 8 34 



384 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORID AE 

elevation of the maxillary in Ph. (?) caffii. Supramaxillary 2 {Smx^, PL 6, fig. 2 ; 
Text-fig. 6) has about the same, almost rectangular shape as in that species but it is 
comparatively longer antero-posteriorly ; its antero-dorsal corner is broken off 
but may have been of about the same shape as in Ph. (?) caffii. The lateral surface 
of supramaxillary 2 shows a horizontal striation, but this striation is by no means as 
dense as in Ph. (?) caffii or in Ph. latiusculus. 

The bones of the infraorbital series are only partly preserved. 

An isolated bone fragment in the snout region looks like an antorhital {Ant?, PI. 6, 
fig. 2), but this interpretation is rather uncertain. 

Another bone fragment dorsal to the anteriormost preserved part of the maxillary 
may belong to infraorbital i {lachrymal) {Ifo-^, PL 6, fig. 2) but it provides no informa- 
tion about the outline of this bone. 

Infraorbital 2 is missing. 

Infraorbital 3 (//03, PL 6, fig. 2 ; Text-fig. 6) is remarkably broad, about twice as 
broad as deep, its postero-dorsal part overlaps the preopercidum and its dorsal 
margin is slightly concave ; its shape is thus quite different from that of the corres- 
ponding bone in Ph. (?) caffii. 

Infraorbital 4 (//04, PL 6, fig. 2 ; Text-fig. 6) is almost complete, nearly rectangular 
and considerably broader than deep. 

Some bone fragments dorsal to infraorbital 4 indicate the presence of an infra- 
orbital 5. 

Between the visible part of the hyomandibular and the anterior margin of the 
operculiun there is a rather large bony plate which is obviously a suborbital {Sbo, 
PL 6, fig. 2 ; Text-fig. 6) ; its margins are, however, only partly visible. 

The preopercuhtm {Pop, PL 6, fig. 2 ; Text-fig. 6) is well preserved, only its upper- 
most end is defective. Its shape is very similar to that of the preoperculum in 
Ph. (?) caffii. A shallow concavity is present ventral to the ill-defined posterior 
angle between the upper and lower limbs of the bone, but there is no real notch as in 
Ph. bechei and Ph. latiusculus. 

The operculum {Op, PL 6, fig. 2 ; Text-fig. 6) seems to be remarkably broad, much 
broader than in the other species treated here, but this may be partly due to pressure 
as the bone is somewhat dislocated. 

The suboperculnm {Sop, PL 6, fig. 2 ; Text-fig. 6), likewise dislocated, is only partly 
preserved ; its postero-dorsal half is missing. 

Of the interoperculum {lop, PL 6, fig. 2 ; Text-fig. 6) only the posterior part is 
exposed ; it seems to have the same general shape as in Ph. (?) caffii, but lacks the 
concentric striations. 



Branchiostegal rays and gular plate 

As in Ph. (?) caffii only some of the posterior branchiostegal rays {R. Br, PL 6, fig. 2) 
are exposed ; they are, however, relatively much longer than in that species. 
The gular plate is not visible. 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHORID AE 385 

Lower jaw 

The lower jaw is exposed to a certain extent, but its anterior part is lacking and 
its dorsal margin is covered by the maxillary bones and cannot be described. It is, 
however, obvious that the lower jaw is relatively much longer and stouter than in 
Ph. (?) caffii. The lateral surface of the dental part of the dentary [De. Spl, PL 6, 
fig. 2 ; Text-fig. 6) is quite smooth, whereas the splenial part, which seems to be 
deeper than in the three species described above, has a rich ganoin ornamentation 
obscuring the tubules of the mandibular sensory line ; the ganoin-covered ridge 
separating these two parts of the dentary is very pronounced. No dentition could 
be observed. 

The angular part of the angulo-supra-angiilar is partly visible but its posterior 
margin is indistinct because of damage ; its lateral surface is ornamented ventrally 
with ganoin patches. 

Sensory canal system of head 

Very little of the sensory canal system of the head can be made out. All the 
bones containing the supraorbital sensory canal are absent and only some parts of 
the infraorbital sensory canal are preserved (Text-fig. 6). In infraorbital 3 the sensory 
canal can be traced but the tubules are difficult to observe ; there are, however, 
traces of at least two rather long, posteriorly directed tubules, and a third, very short 
tubule is given off from the anterior part of the canal. In infraorbital 4 the canal 
itself cannot be seen with accuracy, but a single large, posteriorly directed tubule is 
clearly visible. 

Cephalic division of main lateral line 

Of this part of the sensory canal system only the preopercular sensory canal is 
visible (Text-fig. 6). The canal itself pierces the preoperculum at about the middle 
of its length and, on the whole, lies parallel to its anterior and posterior margins, thus 
not nearer to its posterior margin in the dorsal limb as it is in Ph. (?) caffii and not 
distinctly nearer to its anterior margin as in Ph. bechei and Ph. latiusculus. The 
number of tubules given off from the posterior side of the canal is difficult to deter- 
mine, but two tubides are clearly visible in the dorsal limb of the bone, dorsal to and 
at its posterior angle respectively ; ventral to this angle there is a rather short 
tubule followed more ventrally by about eight wide tubules in the antero-ventral 
third of the bone. Between the second and the third tubules reckoned from above 
there is a structure resembling a postero-dorsally directed tubule or rather, a sensory 
line ; its position posterior to the preopercular sensory line may be an argument 
against the interpretation of it being a supramaxillary line, but as it cannot be 
followed anteriorly because of the damaged condition of the lateral surface of the 
middle part of the preoperculum the question of its real nature must be left open. 
The same is true of the postmaxillary line, the ventralmost part of which may 
perhaps be traced ventral to the damaged middle part of the bone {orp?, PL 6, fig. 2). 
The total number of tubules belonging to the preopercular sensory canal may 



380 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORID AE 

consequently be eleven, perhaps twelve, that is, only a little higher than in Ph. (?) 
caffii ; their arrangement is virtually identical in the two species but those of the 
ventral group are considerably longer, almost reaching the ventral margin of the 
preoperculum ; they are, however, not as long and as numerous as in Ph. bechei 
or Ph. lathtsculus. 

The mandibular sensory canal is not visible and only two tubular openings can be 
traced posteriorly on the splenial part of the dentary. 

A short vertical groove on the lateral surface of the angular is probably an oral 
pit-line {orp, PI. 6, fig. 2). 

Exoskeletal shoulder-girdle and squamation 

No bones belonging to the exoskeletal shoulder-girdle are visible. 

The scales are comparatively thick with even posterior margins. There seem to 
be about 38-40 transverse rows of scales, counted from the posterior margin of the 
operculum to the middle of the caudal fin base, but as the squamation is in part 
rather badly damaged, this count is somewhat uncertain. On the anterior part 
of the body there are some longitudinal rows of scales which are deeper than broad. 
The dorsal margin of each body scale is drawn out into a peg {Sc, PL 6, fig. 2). 



Paired and unpaired fins 

The pectoral, ventral and dorsal fins are missing and only a few lepidotrichia of 
the anal fin are partly visible. 

The caudal fin is hemi-heterocercal ; its dorsal margin carries a dense series of 
rather large fulcra ; along the ventral margin of the caudal fin the fulcra are smaller. 

Remarks. The specimen just described, identified by Woodward (1895) as 
Pholidophorus latiusculus, cannot belong to that species, or to Ph. (?) caffii, as is 
evident from the descriptions given above. On the contrary it seems to me rather 
probable that it may be identical with the second Pholidophorus species originally 
mentioned by Agassiz (1832) from Seefeld, Ph. pusillus, the material of which belonged 
to Dr. Alex. Braun and the Karlsruhe Musernn. Dr. Erwin Jorg, Landessammlungen 
fiir Naturkunde, Karlsruhe, has kindly informed me that no material of Ph. pusillus 
or of Ph. latiusculus now exists in this collection ; it was probably destroyed in 1942. 
The original material of Ph. pusillus must also be considered lost. The identification 
of Ph. pusillus proposed by Kner (1866) refers to some small fish specimens from 
Seefeld ; in the Univ.-Institut fiir Palaontologie und Geologic, Innsbruck, some 
material exists which can be identified as the two upper specimens figured by him on 
pi. 6, fig. 2 (in reality the specimen b on this figure is com^posed of two specimens!). 
Their preservation is, however, rather poor, and they show no details (except some 
branchiostegal rays) which can be directly compared with those preserved in P.4418. 
But there are some facts favouring the identification of this specimen with Ph. 
pusillus. The size and slender shape of the body is about the same and according to 
Kner the number of transverse scale-rows amounts to 38-40, just as I have found in 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORI D AE 387 

the specimen in question. Because of this it seems to me rather improbable that 
two different Pholidophorus species of about the same size and shape should exist 
in the Seefeld deposits. Specimen P.4418 is most probably an unusually well- 
preserved, large specimen of Ph. pusillus. A final decision as to its relationship 
with that species must, however, be postponed until more extensive and better pre- 
served material from Seefeld can be thoroughly investigated. Therefore I have only 
identified the specimen in question as Ph. cf. pusillus Agassiz. 

Horizon and locality. Upper Trias ; Seefeld, Tyrol, Austria. 

Genus PHOLIDOLEPIS nov. 

Preliminary diagnosis. Pholidophoridae of rather small size, as far as known 
similar to Pholidophorus but differing in the following features. Exoskeletal cranial 
bones with very feeble ganoin covering. Supramaxillary 2 with rather well-marked 
process at antero-dorsal corner. Preoperculum with preopercular sensory canal 
running near to anterior margin. Fulcra as far as known absent on all fins except 
along dorsal margin of caudal fin. Scales thin and cycloid, with fine concentric 
striations. 

The generic name Pholidolepis is a composite one derived from Pholidophorus and 
Leptolepis, so as to indicate the probable intermediate position of the new genus 
between the two other genera. 

Type species. Pholidolepis dorsetensis sp. nov. 

Pholidolepis dorsetensis gen. et sp. nov. 

(PL 7 ; Text-figs. 7, 8) 

1895 Pholidophorus caudalis Woodward : 458 [partim). 

Preliminary diagnosis. Pholidolepis of rather small size, up to about no mm. 
in total length. Body elongate, depth about one-fifth of the length without caudal 
(standard length). Head less than one quarter of standard length. Preopercular 
sensory canal with about 15 tubules. 

HoLOTYPE. British Museum (Nat. Hist.) No. 38164. 

Material. In addition to the holotype, Nos. 35725, 38536, P. 3662, P. 44708, 
P.44709 of the British Museum (Natural History) and P. 259, of the D.M.S. Watson 
Collection, Cambridge, have been used for the following description, all more or less 
badly preserved. 

Besides these specimens there is, however, in the British Museum (Natural History) 
a number of still more defective specimens, probably belonging to Ph. dorsetensis : 
Nos. 35562, 38537, 39862, 43007, P.939, P939&, P.939C, P4370. and P.44707. 

Description. The holotype has a total length of about no mm. ; length from 
tip of snout to posterior margins of hypurals (standard length) about 95 mm. 
Specimen P. 44708 has a standard length of about 98 mm., P. 3662 of about 80 mm., 
which lengths constitute the range for the specimens investigated. 



388 CERTAIN TRIASSIC AND LIASSIC PH OLI DOPH ORI D AE 

The body is decidedly more elongate than in Ph. caudalis, to which species the 
specimens in question have been referred by Woodward, but as all specimens are 
crushed the true depth of the body cannot be given with accuracy ; it may be esti- 
mated as about one-fifth of the standard length. The length of the skull seems to be 
less than one quarter of the standard length. 

The dermal bones of the head are comparatively thin ; as far as can be seen only 
the bones of the jaws show a surface ornamentation. 



Exoskeletal skull roof 

Only very little can be made out regarding the bones belonging to the exoskeletal 
skull roof because of the bad state of preservation. 

The premaxillary is not visible in any of the specimens investigated. 

The rostral may be present in Watson Colin. No. P. 259, but only in a defective 
state. 

The nasal (Na, PL 7, fig. 3 ; Text-figs. 7, 8) is rather well preserved in the holotype. 
It is an elongate, comparatively large bone reminiscent of the corresponding bone in 
Ph. hechei and situated lateral to the foremost part of the lateral margin of the frontal ; 




Fig. 7. Pholidolepis dorsetensis gen. et sp. nov. Attempted restoration of head in lateral view. 
x6-3. 

Ang, angular ; De. Spl, dentary ; Fr, frontal ; lop, interoperculum ; Mx, maxillary ; Na, 
nasal ; Op, operculum ; Pop, preoperculum ; Smx^, Smx2, anterior and posterior supra- 
maxillaries ; Sop, suboperculum ; poc, preopercular sensory canal ; soc, supraorbital sensory 
canal. 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORIDAE 389 

anteriorly it is rounded and reaches about as far forwards as the anterior tip of the 
frontal. Its lateral outline cannot, however, be determined exactly because of 
damage and consequently it is impossible to say whether a notch or fenestra for the 
posterior nostril is present as in Ph. bechei. 

The frontal {Fr, PL 7, figs. 1,3; Text-figs. 7, 8) is preserved in the holotype, in 
38536 and in Watson Colin. No. P. 259, but in all these specimens it is more or less 
defective because of crushing. Its anterior half is, judging from the holotype, rather 
broad but tapers towards the anterior rounded tip at the level of the nasal. The 
posterior half broadens considerably but its posterior margin cannot be accurately 
defined in any specimens. There are no traces of surface ornamentation on the frontal. 

No supraorbitals can be seen clearly in any specimens but some small bone frag- 
ments lateral to the lateral margins of the frontal and nasal in the holotype {So, 
PI. 7, fig. 3) may belong to an anterior supraorbital similar to that bone in Ph. bechei. 

Only the sensory canal of the dermosphenotic seems to be present in the holotype, 
the rest of the bone is missing. In the other specimens investigated this bone is 
not visible. 

The parietal, the dermopterotic and the extrascapular are, in all specimens, too 
damaged for description. 

Dermal bones of cheek and opercular apparatus 

The maxillary {Mx, PL 7, figs. 1-3 ; Text-fig. 7) is more or less completely preserved 
in the holotype as well as in 38536 and P.44708, and Watson Colin. No. P. 259, but in 
none of them is it undamaged and its anterior, antero-mesially directed tip is not 
visible. Its posteriorly directed, lateral part is, as usual, evenly curved with the 
convexity directed ventrally. Its lateral surface is provided with a weak ornamenta- 
tion, consisting of more or less oblique striations as can be seen in the holotype and 
particularly in P.44708 (PL 7, fig. 2) . In the latter the ventral border of the maxillary 
carries a dentition similar to that in Leptolepis. 

The supramaxillaries are only visible in P.44708 and Watson Colin. No. P. 259. 
The anterior one, supramaxillary i {Smx-^^, PL 7, fig. 2), is partly preserved in the 
first named specimen and seems to be much longer antero-posteriorly than the 
corresponding bone in Pholidophorus , thus more like that of Leptolepis, but as its 
anterior part is damaged its outline cannot be made out with accuracy. If there 
exists any surface ornamentation on supramaxillary i it must be only weakly 
developed. Supramaxillary 2 {Smx^, PL 7, fig. 2) is also best preserved in P.44708 ; 
its antero-dorsal process is decidedly more pronounced than in Pholidophorus but 
not nearly so long as in Leptolepis. The lateral surface of supramaxillary 2 carries a 
weak longitudinal striation somewhat recalling that in Ph. latiusculus but much 
weaker. Both supramaxillaries seem to overlap the dorsal margin of the maxillary 
but only to a small degree. 

The bones of the infraorbital series as well as the suborbital are practically missing 
in all specimens. 

The preoperculum {Pop, PL 7, figs. 1,3; Text -fig. 7) is present in most specimens 
but in none is it undamaged. It is best preserved in 38536 and has the same general 



390 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPH OR I D AE 




Fig. 8. Pholidolepis dorsetensis gen. et sp. nov. Attempted restoration of head 
in dorsal view, x 6'3. Lettering as in Fig. 7. 

shape as in Leptolepis, the anterior margins of the dorsal and the antero-ventral 
hmbs meeting to enclose an angle, but it has a deep notch on its posterior margin as 
in Ph. bechei. There is no process on its anterior margin at the angle as there is, for 
example, in Leptolepis normandica. 

The operculum [Op, Text-fig. 7) is very damaged by crushing or is lacking in all 
specimens except Watson Colin. No. P. 259. Judging from this specimen its general 
shape is about the same as in Pholidophorus. 

The suhoperculum {Sop, Text-fig. 7) is also only preserved in Watson Colin. No. 
P. 259. It seems to be comparatively deep, and the process at its antero-dorsal 
corner is longer and more pointed than in Ph. bechei. 

Only the posterior part of the interoperctilum {lop, Text-fig. 7) is exposed in the 
specimen just named and in 38536, and nothing can be said about its general shape. 



Lower jaw 

The lower jaw is partly visible in the holotype and in 38536 and P. 44708, and 
Watson Colin. No. P. 259. Its outhne cannot be determined, but in all specimens 
mentioned the anterior part of the dentary {De. Spl, PI. 7, figs. 1-3 ; Text-fig. 7) 
ascends rather gently as in Pholidophorus, not abruptly as in Leptolepis ; from this 
it may be concluded that the general shape of the lower jaw is also almost as in 
Pholidophorus. The ridge separating the dental and the comparatively low splenial 
parts of the dentary is clearly visible. In specimen P. 44708 (PI. 7, fig. 2) the lateral 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPHORI D AE 391 

surface of the angular {Ang, PL 7, fig. 2) is provided with a surface ornamentation 
of thin longitudinal ridges or streaks similar to those on the maxillary. 
A gidar plate is not visible in any of the specimens investigated. 

Sensory canal system of head 

_ 'le sensory canal system of the head is only partly visible and the description 
must therefore be rather incomplete. 

Fart of the supraorbital sensory canal {soc, PL 7, figs, i, 3 ; Text-figs. 7, 8) can 
be seen in the holotype, in 38536 and in Watson Colin. No. P. 259. In its foremost 
part, piercing the nasal bone, the canal is straight and continues straight backwards 
in the anterior, narrower part of the frontal. As the frontal broadens the canal bends 
laterally and then runs obliquely postero-mesially ; its posterior part in the parietal 
cannot be followed. No tubules or pores can be observed with accuracy except in 
Watson Colin. No. P. 259, where three postero-mesially directed tubules are given off 
from the mesial side just before and at the bend of the canal. 

Almost nothing can be made out regarding the infraorbital sensory canal. In 
the holotype only a small part of a sensory canal is present, probably the canal in the 
dermosphenotic {Dsph, PL 7, fig. 3), with the backward bend of the main canal and 
the antero-dorsally directed tubule. 

Cephalic division of main lateral line 

The cephalic division of the main lateral line can only be followed in the pre- 
operculum. 

The preopercular sensory canal {poc, PL 7, fig. 3) and its tubules follow the same 
general pattern as in Ph. bechei, but the number of tubules is lower. In the ventral 
limb of the preoperculum there are in 38536 eleven tubiiles, obviously corresponding 
to the 13 ventrally situated tubules in Ph. bechei ; the posterior ones are, as in that 
species, long, narrow and curved with the convexity anteriorly directed. Dorsal to 
these tubules there are four shorter tubules, corresponding to the five to six shorter 
tubules in Ph. bechei. The total number of tubules is thus 15 in this specimen ; 
in the others investigated the preoperculum is too damaged to allow a count of the 
tubules, but as far as can be observed the general pattern is the same as that just 
described for 38536. 

Exoskeletal shoulder girdle and squamation 

Parts of the cleithrum are exposed in many of the specimens but in no case can it be 
studied in any detail. No other bones belonging to the exoskeletal shoulder-girdle 
can be identified. 

The squamation consists of thin cycloid scales with delicate concentric striations. 

The lateral line 
No lateral line can be seen with certainty in any of the specimens available. 



392 CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORIDAE 

A xial skeleton and paired and unpaired fins 

The vertebral centra are thin bony cyhnders which are, however, crushed or dis- 
placed in all specimens investigated ; their undamaged shape cannot be described 
nor can their number be determined with any degree of accuracy. The neural and 
haemal spines are comparatively long. 

The fins are in part rather well preserved but in the dorsal and anal fins the lepido- 
trichia are poorly preserved. No fulcra can be observed on the pectoral, ventral, 
dorsal or anal fins, but with regard to the two last named fins this statement needs 
confirmation on better preserved material. 

The pectoral fm has a rather high number of lepidotrichia ; in P. 3662 and P.44709, 
and Watson Colin. No. P. 259 I have coimted 19 rays, in P.44708 20 rays. 

In the ventral fin the number of lepidotrichia is also rather high ; in P.44708 and 
P. 44709 and Watson Colin. No. P. 259 their number is 15 or about as in Ph. bechei. 

The dorsal fin is poorly preserved in all specimens, but in P. 44709 the number of 
dorsal radials can be determined as 12, the anterior one as usual being composed of 
two elements. 

The anal fm has 8 radials in the holotype and in 38536 and P.44709. 

The caudal fin carries along its dorsal margin rather long and stout fulcra ; no 
fulcra have been observed on the ventral margin of the fin. 

Remarks. There is no doubt that the specimens just described were erroneously 
identified as Pholidophorus caudalis Woodward, as can be seen from a comparison 
with the following description (see p. 411), and I have been unable to find another 
described species to which they can be referred. Consequently I have assigned them 
to a new species for which I propose the name Pholidolepis dorsetensis. Unfor- 
tunately the material on which the new species is based is rather defective, and I am 
not fully convinced that all specimens here referred to Ph. dorsetensis really belong 
to that species. In particidar I am in doubt regarding Watson Colin. No. P. 259, 
which shows very few details directly comparable with those exposed in the other 
specimens ; it is especially unfortunate that almost nothing of the characteristic 
preopercidum is preserved. On the other hand I have not found any differences 
justifying its separation from the other specimens. More complete and better 
preserved material is necessary for a fuller description of Ph. dorsetensis. 

Horizon and locality. Lower Lias ; Lyme Regis, Dorset. 

Genus PHOLIDOPHOROIDES Woodward 

1941 Pholidophoroides Woodward : 89. 

Emended diagnosis. Pholidophoridae of medium size. Exoskeletal cranial 
bones and scales with ganoin covering. Nasal well developed and well separated 
from its antimere by the frontals, anteriorly reaching level of anterior margin of 
frontals. A single, well-developed supraorbital. Maxillary rather stout and deep 
with more or less pronounced convexity on dorsal margin ventral to infraorbital i 
(lachrymal) and supramaxillary i ; posterior margin notched. Two supramaxil- 
laries, not markedly overlapping dorsal margin of maxillary, supramaxillary 2 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORIDAE 393 

without marked process at antero-dorsal corner. Antorbital well developed, more 
than five (seven?) infraorbitals. Preoperculum with preopercular sensory canal 
running along middle of bone, about equidistant from anterior and posterior margins. 
Lower jaw not markedly deep, greatest depth in posterior third of its length ; dentary 
with smooth dental part, separated from splenial part by strong ridge. Dorsal 
fin above interspace between ventral and anal fins. Fulcra present along anterior 
margin of paired as well as unpaired fins. Caudal fin hemi-heterocercal with dorsal 
and ventral margins fulcrated. Scales moderately thick ; some longitudinal series 
of flank-scales higher than broad. 

Type species. Pholidophoroides crenulata (Egerton). 

Remarks. When creating the new genus Pholidophoroides Woodward (1941 : 89) 
diagnosed it in the following manner : " One species from the Lower Lias, P. crenu- 
latus Egerton, is distinguished from Pholidophorus by its smaller scales, which are 
less deepened on the flank. The scales are arranged in from 50 to 60 transverse 
rows, and those of the anterior part of the lateral line are less than twice as deep as 
broad. The scales are also comparatively thin, often displaying their concentric 
lines of growth. The fins are small like those of Pholidophorus. The maxilla seems 
to be comparatively stout. Ring vertebrae are conspicuous in the abdominal region. 

P. crenulatus may therefore be regarded as the type-species of a distinct genus, 
Pholidophoroides. . . . The Lower Liassic species, P. caudalis A. S. Woodw., probably 
also belongs to Pholidophoroides." 

The diagnosis given above is mainly based on the taxonomic characteristics of the 
redescribed type species, but also taking into consideration the other species here 
ascribed to the genus Pholidophoroides. 

Pholidophoroides crenulata (Egerton) 
(Pis. 8, 9, 10 ; PI. 15, figs. 3, 4, 5, 8 ; Text-figs. 9-11) 

1843 Pholidophorus crenulatus Egerton : 184. 

1852 Pholidophorus crenulatus Egerton ; Egerton, pi. 5. 

1887 Isopholis crenulatus (Egerton) Zittel : 216. 

1890 Pholidophorus crenulatus Egerton ; Woodward & Sherborn : 146. 

1895 Pholidophorus crenulatus Egerton ; Woodward : 463, pi. 12, fig. 6. 

1941 Pholidophoroides crenulatus (Egerton) Woodward : 89. 

Diagnosis. Pholidophoroides of medium size, at least up to 150 mm. in total 
length. Greatest depth of the body about one quarter of the standard length or 
slightly more {ca. 25-30% of that length). Depth of the caudal peduncle less than 
one-tenth [ca. 8-9%) of the standard length. Maxillary deep. Preopercular sensory 
canal with 16 or 17 tubules. Scales rather thin, in about 45-50 transverse rows 
reckoned from the hind margin of operculum to the middle of the caudal fin ; about 
four longitudinal rows of body scales somewhat deeper than broad and with the 
hind margin crenulated. 

Lectotype. British Museimi (Natural History) Nos. P. 572 and P. 573 are the 
two specimens originally described and figured by Egerton (1843, 1852) and designated 



394 CERTAIN TRIASSIC AND LIASSIC PH OLI D OPH O RI D AE 

as " The two type specimens " by Woodward (1895 : 463). P. 573 is the more com- 
plete specimen (standard length about 82 mm., height of body about 21 mm., height 
of caudal pedimcle 7-5 mm.) and is therefore selected as lectotype. 

Material. In addition to the lectotype, the specimens mainly used for the 
following description are P. 153, P.421, P. 572, P. 1046a, b, P. 4415, 38110, 38739, 
38738 ; they agree in all comparable features with P. 573. 

Description. The total length of the largest specimen, 'P. 1046a, is 150 mm. ; 
its length from the tip of the snout to the hindmost scale in the lateral line (standard 
length) is about 125 mm. The other specimens in the British Museum (Natural 
History) have a standard length from 120 to 74 mm. The greatest depth of the body 
is about one quarter of the standard length or slightly more [ca. 25-30% of this 
length) ; the depth of the caudal peduncle is somewhat less than one-tenth (8-9%) 
of the standard length. The length of the head is about one quarter of the standard 
length, generally slightly less {ca. 22-25%). 

The ventral fins are situated approximately midway between the tip of the snout 
and the hindmost lateral line scale and the distance between the base of the ventrals 
and the origin of the anal is about 75-85% of the distance between the bases of the 
pectoral and ventral fins. 

A good figure of the entire fish is given by Woodward (1895, pi. 12, fig. 6). 

The specimens most thoroughly investigated are rather well preserved with the 
bones of the skull exposed to a great extent, and consequently they allow the follow- 
ing description to be made. 

Exoskeletal skull roof 

The premaxillary {Pmx, PL 10, figs, i, 2 ; Text-fig. 9) is visible in P. 1046a, b, 
and to some extent also in P.4415 and 38738. It is comparatively stout with the 
postero-dorsal margin weakly S-shaped. The straight antero-ventral margin is 
serrated, forming a single row of teeth. The lateral surface of the premaxillary is 
densely set with ganoin tuberculations. 

The rostral {Ro, PL 9 ; Text-figs. 9, 10) is best preserved in 381 10, but cannot be 
studied in any detail. The part pierced by the ethmoidal commissure is a little 
swollen dorsally ; posteriorly its lateral parts are concave and rather thin at the 
margin. 

The nasal (Na, Pis. 8, 9 ; Text-figs. 9, 10) is also best exposed in 38110. It is an 
oblong, rather broad bone, situated posterior to the lateral parts of the rostral. 
It is, on the whole, oval but its antero-lateral and postero-lateral margins (lateral 
to the posterior opening for the supraorbital sensory canal) are concave, probably 
marking the postero-mesial and antero-mesial limits of the anterior and posterior 
nasal openings respectively. The nasals are separated by the narrow anterior part 
of the frontals. 

The frontal (Fr, Pis. 8, 9 ; PL 10, fig. 2 ; PL 15, fig. 4 ; Text-figs. 9, 10) is rather 
narrow in its anterior part, mesial to the nasal, but it broadens considerably posterior 
to the nasal ; from there its lateral margin runs at first almost straight posteriorly, 
but behind the orbit the bone widens again and has its greatest breadth in its posterior 



CERTAIN TRIASSIC AND LIASSIC PHOLI DOPH O R I D A E 



395 



Dpt Ext 

''( P^a 1 sxom I 

1 I 



I 



1^ 
Ro \ 

\ Ant 



Ljc.com \ 



Pmx- 




FiG. 9. Pholidophoroides crenulata (Egerton). Attempted restoration of head in lateral view. 
X 5 approx. 

Ang, angular ; Ant, antorbital ; De. Spl, dentary ; Dpt, dermopterotic ; Dsph, dermo- 
sphenotic ; Ext, extrascapular ; Fr, frontal ; Ifo^-Ifo-,, infraorbitals i to 7 ; lop, interoper- 
culum ; Mx, maxillary ; Na, nasal ; Op, operculum ; Pa, parietal ; Pmx, premaxillary ; 
Pop, preoperculum ; Ro, rostral; Sbo, suborbital; " Sbo", " accessory suborbital " ; Scl, 
supracleithrum ; Smx^, Smx^, anterior and posterior supramaxillaries ; So, supraorbital ; 
Sop, suboperculum ; Ssc, suprascapula ; ant. br, antorbital branch of infraorbital sensory 
canal ; ap, anterior pit-line ; ifc, infraorbital sensory canal ; ifc. com, ethmoidal commissure ; 
mdc, mandibular sensory canal ; mp, middle pit-line ; poc. preopercular sensory canal ; pp, 
posterior pit-line ; s. com, supratemporal commissure ; soc, supraorbital sensory canal. 



part. Its postero-lateral corner is rounded off and its posterior margin is irregularly 
wavy. The suture between the frontals is practically straight anteriorly ; posteriorly 
its course is difficult to follow, but it does not seem to be markedly sinuous. 

A strong supraorbital {So, PL 9 ; Text -figs. 9, 10) is situated behind the posterior 
nasal opening and lateral to the broadened part of the frontal posterior to the nasal. 
Anteriorly its dorsal surface bulges a little and is ornamented with some small 
elevations. In dorsal aspect the lateral border of the supraorbital is almost straight, 
ending anteriorly in a point. 

A very small bone between the lateral margin of the frontal and the anterior tip 
of the dermosphenotic in 38739 {So?, PL 8 ; PL 15, fig. 4) may perhaps be a small 
second supraorbital ; in P.1046& a similar small bone may be traced, but as I have 



396 CERTAIN TRIASSIC AND LIASSIC PHOL IDOPHORI D AE 

not found a corresponding element in any of the other specimens investigated, it 
may perhaps be only a fragment of the anterior part of the dermosphenotic or of the 
lateral frontal margin. 

The dermosphenotic {Dsph, Pis. 8, 9 ; PI. 10, fig. 2 ; PI. 15, fig. 4 ; Text-figs. 9, 10) 
is mostly more or less crushed on the underlying autosphenotic. Its anterior part, 
situated along the postero-lateral margin of the frontal, is pointed and a little curved 
and constitutes the postero-dorsal border of the orbit. Its posterior margin cannot 
be determined with accuracy, but according to P. 1046ft it seems to be rounded off. 

The margins of the parietal {Pa, PI. 10, fig. 2 ; PI. 15, fig. 4 ; Text-figs. 9, 10) are 
difficult to determine, as the skull roof of the single specimen showing the cranial 
bones in dorsal view, 38110, is much ciushed in its posterior part, but judging from 
38739 it has a rather wide extension laterally, thus being much broader than long. 

The dermopterotic {Dpt, Pis. 8, 9 ; PI. 10, fig. 2 ; PI. 15, fig. 4 ; Text-figs. 9, 10) 
is comparatively narrow in its anterior part between the frontal and the dermo- 
sphenotic but broadens continuously backwards, attaining its greatest breadth at 
its straight posterior margin. 

The shape of the extrascapular [Ext, PI. 9 ; PL 10, fig. 2 ; Text-figs. 9, 10) cannot 
be determined ; its lateral part is, however, almost triangular with the postero- 
lateral margin a little convex and the postero-mesial margin slightly concave. The 
mesial part of the extrascapular is not preserved in any of the specimens investigated. 

Dermal bones of cheek and opercular apparatus 

The maxillary {Mx, Pis. 8, 9 ; PI. 10, figs, i, 2 ; Text-fig. 9) is strikingly large and 
stout. Its anterior, mesially directed part is only partly visible in the specimens 
investigated, but seems to be rather low ; its dorsal margin has an arched, thickened 
edge with the convexity directed ventrally, fitting the concavity on the postero-dorsal 
margin of the premaxillary ; the surface covered by the premaxillary is smooth. 
The lateral, posteriorly directed part of the maxillary increases rapidly in height ; 
its ventral margin is evenly convex, but its dorsal margin, on the whole concave, 
has a marked convexity below infraorbital i and supramaxillary i. The posterior 
margin of the maxillary is rather deeply notched. The anterior half of the lateral 
surface has, nearest to the premaxillary, an ornamentation of densely set tubercula- 
tions like those on the premaxillary ; posteriorly the ornamentation becomes a 
longitudinal striation parallel to the margins. The posterior half of the lateral 
margin is smooth except for some longitudinal striations along the dorsal margin ; 
in P.4415 there are, also, some longitudinal ganoin streaks parallel to the ventral 
margin. A delicate comb-like dentition, about as in Leptolepis, is present along 
the posterior half of the ventral margin, the teeth decreasing in length anteriorly. 
No teeth can be observed along the anterior half of the ventral margin, but whether 
this is the actual condition or only dependent upon the state of preservation, must 
remain undecided. 

Two supramaxillaries are situated dorsal to the maxillary, not markedly over- 
lapping its dorsal margin. The anterior one, supramaxillary i (5m;tj, Pis. 8, g ; 
PI. 10, figs. I, 2 ; Text-fig. 9), is rather small, almost semicircular, and has a well- 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORI D AE 

ifc. corn 



397 




Fig. io. Pholidophoroides crenulata (Egerton). Attempted restoration of head in dorsal view, 
the snout flattened, with (right) the probable mutual positions of the bones of the snout. 
X 5 approx. Lettering as in Fig. g. 

marked concentric striation on its lateral surface. The posterior one, supramaxillary 
2 {Smx^, PI. 8 ; PL lo, figs, i, 2 ; Text-fig. 9), is considerably larger ; its anterior 
margin is concave, fitting the convexity of supramaxillary i, but its antero-dorsal 
corner is not markedly produced into a process. The dorsal and ventral margins of 
supramaxillary 2 are evenly convergent towards the posterior tip of the bone. 
Supramaxillary 2 also has a well-marked concentric striation on its lateral surface. 

The bones of the infraorbital series are generally well preserved, only infraorbital 2 
is lacking in all specimens investigated. 

The antorbital {Ant, Pis. 8, 9 ; PI. 10, fig. 2 ; Text-figs. 9, 10) is an almost J.-shaped 
bone situated ventro-lateral to the rostral and the nasal ; its posterior tip reaches 
the antero-lateral tip of the supraorbital. 

Ventral to the antorbital and between it and the anterior part of the maxillary and 
supramaxillary i there is a large infraorbital i or lachrymal {Ifo-^^, PI. 8 ; PL 10, 
figs. I, 2 ; Text-fig. 9). Its anterior margin is semicircular and its ventral margin is 
slightly convex ; its dorsal margin is not clearly visible but is probably slightly 
concave. 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORI D AE 



Infraorbital 3 (//og, Pis. 8, 9 ; PL 10, figs, i, 2 ; Text-fig. 9) is comparatively large, 
stretching from the orbit to the anterior margin of the preoperculum ; its antero- 
ventral margin is slightly concave. 

The following bones in the series, dorsal to infraorbital 3, are crushed, displaced 
or lacking in most specimens, but judging from 38110 and P. 1046a and b there are 
about four small, square infraorbitals between infraorbital 3 and the dermosphenotic 
(//04, //05, //Og, //07, PI. 9 ; Text-figs. 9, 11) ; in 38110 there seems to exist one more 
element in the series on the right side (//Og?, PI. 9 ; Text-fig. 11), but here the arrange- 
ment is rather obscure. 



--y -Dsph 





Fig. II. Pholidophoroides crenulata (Egerton). Infraorbital 3, postorbital infraorbitals and 
suborbitals (same specimen as on PI. 9). Lettering as in Fig. 9 ; A right side, B left side. B.M. 
38110. X 5 approx. 

A large suborbital [Sbo, Pis. 8, 9 ; PL 10, fig. 2 ; Text-fig. 9) bone is situated 
posterior to infraorbitals 4-7, covering the hyomandibular ; as a rule it is crushed 
in the middle on to the strong lateral crest of this latter bone. Dorsal to the sub- 
orbital there are in 38110 (" Sbo " , PL 9 ; Text-figs. 9, 11) two small bones on the 
right side and a single similar bone on the left side, tapering posteriorly and, like the 
suborbital, with a conspicuous concentric surface striation. A single similar bone is 
also seen in 38738. It is difficult to interpret these bones ; they may be homologous 
to the supraspiracular plates described by Stensio (1932, text-fig. 67) in Perleidus 
stoschiensis Stensio and by Lehman (1952, text-figs. 85, 86) in P. madagascariensis 
Piveteau. But they are more likely to be " accessory suborbitals ". In many 
holosteans there is a series of suborbitals posterior to the infraorbitals which may 
fuse or remain separate in various ways. Thus, for example the posterior part of 
infraorbital 3 in Pholidophoroides as well as in Leptolepis is probably the originally 
ventralmost suborbital, secondarily fused with infraorbital 3 (c/. Stensio 1947 : 164), 
whereas the others as a rule fuse into a single, large suborbital. In Ph. crenulata the 
dorsalmost component or components may remain free, at least in some specimens. 

The peculiar condition of the dorsalmost infraorbitals and the " accessory sub- 
orbitals " on the right side in 38110 is also difficult to interpret. It may be due to 
an ontogenetic disturbance or perhaps to regeneration after an injury. 



CERTAIN TRIASSIC AND LIASSIC PHOLIDOPHORI D AE 399 

The preoperculum (Pop, Pis. 8, 9 ; PL 10, figs, i, 2 ; Text-fig. 9) is a weakly 
crescentic bone with a dorsal, more or less vertical limb and an antero-ventrally 
directed ventral limb, much broader than the dorsal one. Its posterior margin has, 
ventral to the more or less pronounced angle between the two limbs, a characteristic 
shallow concavity or indentation. 

The operculum {Op, Pis. 8,9; PL 10, figs. 1,2; Text-fig. 9) is roughly triangular ; 
its anterior, vertical margin and its postero-ventral margin are straight and its 
dorsal margin rounded off. The anterior margin is thickened ventral to the facet 
for the processus opercularis of the hyomandibular. 

The suboperculum {Sop, Pis. 8, 9 ; PL 10, figs, i, 2 ; Text-fig. 9) is comparatively 
large and its antero-dorsal process anterior to the ventral tip of the operculum is 
remarkably broad and strong. According to 38739, P. 10466 and P. 4415 the posterior 
margin of the suboperculum has dorsally a shallow concavity or indentation. 

The interoperculum {lop, PL 8 ; PL 10, figs, i, 2 ; Text-fig. 9) has a roughly 
triangular shape, with the posterior and ventral margins a little concave. 

Branchiostegal rays and gular plate 

There seems to be a rather large number of branchiostegal rays ; they are best 
exposed in P.4415 {R. Br, PL 10, fig. i), on which 16 rays can be seen, but more small 
ones may exist in front of the anteriormost of the exposed rays. 

P.10466 shows parts of a rather large median gular plate {Gu, PL 10, fig. 2). Its 
anterior tip is a little bent upwards, but as the lateral margin is broken off, nothing 
can be stated regarding its shape. 

Lower jaw 

The lower jaw is generally only partly exposed, its dorsal margin being covered by 
other bones in all specimens investigated except in P. 153 (PL 15, fig. 5), where the 
lower jaw is isolated and its outline can be determined rather exactly ; its greatest 
depth is somewhat less than half of its length. 

The ventral, splenial part of the dentary {De. Spl, PL 8 ; PL 10, figs. 1,2; PL 15, 
fig. 5 ; Text -fig. 9) is limited dorsally by a well-marked ridge, and its lateral surface 
is ornamented with longitudinal ganoin striations and rugosities. The